neumania papillator is a water mite in the genus neumania and is known in behavioural ecology as an example of sensory exploitation - males of this species hijack existing female sensory abilities for their own gain in courtship .

neumania papillator

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discord between field and laboratory sex ratios of the water mite neumania papillator marshall ( acari : unionicolidae ) .
the sensory exploitation hypothesis explains male secondary sexual characters as adaptations to exploit female responses that evolved in non - sexual contexts . the water mite neumania papillator takes up a characteristic posture , the net - stance , in order to detect vibrations produced by copepod prey . courting males vibrate their legs ( ‘male courtship trembling’ ) near females , and previous . . . [ show full abstract ]
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present address : department of biological sciences , university of calgary , calgary , alberta , t2n 1n4 canada .
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to cite this page : myers , p . , r . espinosa , c . s . parr , t . jones , g . s . hammond , and t . a . dewey . 2018 . the animal diversity web ( online ) . accessed at https : / / animaldiversity . org .
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. . . pre - existing biases in the receiver ' s sensory ( sen - sory exploitation ; ryan et al . , 1990 ) or cognitive ( sensory trap ; christy , 1995 ) systems can direct the evolution of communication systems ( ryan & cummings , 2013 ) . for example , female responses to food cues are exploited by males in many taxa ( proctor , 1991 ; rodd et al . , 2002 ; garcia & ramirez , 2005 ; mart ın & l opez , 2008 ; kolm et al . , 2012 ) . receiver biases are generally supported by evidence that ( i ) the response is shaped by natural selection rather than sexual selection , and ( ii ) the response evolved prior to the signal . . . .
. . . the hypothesis that a receiver bias shaped the evolu - tion of 3kpzs communication is supported by the exist - ing data but should nonetheless be considered bearing some scepticism . the most conclusive lines of evidence for a receiver bias are ( i ) a demonstrated response to the trait formed under natural selection rather than sexual selection , and ( ii ) a phylogenetic reconstruction that indicates the preference arose before the male sig - nal ( proctor , 1991 ) . the evidence that larval lamprey releases 3kpzs and adult lamprey orient towards 3kpzs during migration offers strong support that female lam - preys respond to 3kpzs in a nonsexual context ( buchinger et al . , 2013 ; brant et al . , 2016 ) . . . .
. . . finally , in addition to trait quality , the short - term condition of an individual also influences mate choice in invertebrates . for instance , hungry females mate less selectively and often more frequently than well - fed females in a number of insects ( proctor , 1991 ; poulin , 1994 ; simmons , 1994 ) . copepods likely experience instances of food limitation in the field ( bergreen et al . , 1988 ; durbin et al . , 1992 ; müller - navarra and lampert , 1996 ; gulati and demott , 1997 ; sterner and schulz , 1998 ) . . . .
. . . the size of water mite larvae from hygrobates , unionicola and piona genera after hatching is ca . 200 µm ( wainstein 1980 , proctor 1991 tuzovsky 2011 ) . in our study , the sizes of larval hygrobates sp . . . .
. . . when a sensory system has evolved due to selection in one context , it may allow organisms to exploit signals and cues for other purposes . studies investigating this pre - existing sensory bias hypothesis have focused on signals and cues that are incorporated from other contexts into mating ( proctor , 1991 ; weller , jacobson , & conner , 1999 ) . less attention , however , has been given to how sensory systems that evolve for mating may guide the use of signals or cues to avoid predators , or to forage . . . .
. . . finally , in addition to trait quality , the short - term condition of an individual also influences mate choice in invertebrates . for instance , hungry females mate less selectively and often more frequently than well - fed females in a number of insects ( proctor , 1991 ; poulin , 1994 ; simmons , 1994 ) . copepods likely experience instances of food limitation in the field ( durbin et al . , 1992 ; bergreen et al . , 1988 ; müller - navarra and lampert , 1996 ; gulati and demott , 1997 ; sterner and schulz , 1998 ) . . . .
. . . irwin , 2000 ) . our results also parallel research that shows how mating signals can adapt to pre - existing biases for particular types of sensory information ( proctor , 1991 ) . however , in this study we provide an example of a different type of contextual shift , namely from a predator context , and potentially an antipredator function , to a sexual display context . . . .
. . . stålhandske ( 2002 ) found that males with white wrapped gifts are accepted faster than males with dark gifts and suggested males exploit the female maternal care instinct , but recent studies rejected this idea and suggested male exploitation of female foraging motivation ( bilde et al . 2007 ; prokop and maxwell 2009 ) . thus , females would respond to conspicuous male traits because these traits stimulate their sensory system ( westeberhard 1984 ; basolo 1991 ) , an idea suggested in several species ( ryan et al . 1990 ; basolo 1990 ; proctor 1991 proctor , 1992 christy 1995 ; sakaluk 1984 sakaluk , 2000 rodd et al . 2002 ; christy et al . 2003a ; b ; madden and tanner 2003 ; reinhardt et al . 2009 ) . nevertheless , p . mirabilis males in poor condition are usually limited in their ability to construct gifts and perform courtship , experiencing lower reproductive success than males in good condition ( albo et al . 2011b ; albo et al . 2012 ) . . . .
. . . some studies offer clear evidence of se as a true alternative to costly signaling ( ce ) . for example , in a well - documented case , male water mites mimic prey in order to attract the attention of females ( proctor 1991 proctor , 1992 ) . this case illustrates the strong version of se because it precludes cs to operate . . . .
sensory exploitation and the evolution of male mating behaviour : a cladistic test using water mites . . .
although copulation is a widespread behavior with multiple origins , hypotheses about selective forces behind its evolution have not been tested by the comparative method . because copulation has arisen many times in the water mites ( acari : parasitengona ) , they are good subjects for a comparative study of copulation . i determined that copulation evolved 91 times in the 343 extant genera . there . . . [ show full abstract ]
chemoreception in four species of water mites ( acari : hydrachnida ) : behavioural and morphological ev . . .
silver nitrate staining of the solenidial and eupathidial sensilla on the palpi , tarsi and tibiae of legs i and ii of arrenurus acutus , geayia ovata , mideopsis reelfootensis and albia caerulea , along with ultrastructural studies , showed that these setae have a porous shaft cuticle . these setae are hollow and contain dendrites in their lumens and more than one neuron is associated with each . . . [ show full abstract ]
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received 2007 oct 24 ; revised 2007 nov 15 ; accepted 2007 nov 16 .
arthropod filiform hairs detect air ( or water ) particle displacements ( görner & andrews 1969 ) and are among the most sensitive animal sensory organs ( shimozawa et al . 2003 ) . such near - field particle movements are used in intraspecific communication by a restricted group of insects , but these use specialized organs for particle movement detection , such as the johnston ' s organ of some insect antennae ( e . g . johnston 1855 ; tauber & eberl 2003 ; gibson & russell 2006 ; tsujiuchi et al . 2007 ) . whether filiform hairs also receive such intraspecific signals is unclear , but given their abundance across different arthropod taxa , their use in signal reception may indicate that near - field sounds are a far more common communicatory modality than previously realized .
here we investigate whether arachnid trichobothria function in intraspecific near - field sound detection during the agonistic contests of whip spiders ( arachnida , amblypygi ) . whip spiders encompass their own arachnid order and possess enlarged , spiny pedipalps , and modified ( antenniform ) first legs used as feelers rather than walking legs . contests consist of a sequence of ritualized displays , the most striking of which is a rapid vibration of the displayer ' s outstretched antenniform leg close to its opponent ( alexander 1962 ; weygoldt 2000 ; fowler - finn & hebets 2006 ) . in complete darkness ( excluding visual detection ) , this display is performed for longer by contest winners than losers , and probably conveys information used to resolve contest outcome ( fowler - finn & hebets 2006 ) .
we show that the agonistic antenniform leg vibrations ( alvs ) of the whip spider phrynus marginemaculatus are received by trichobothria and are suited to eliciting a particularly strong response in them . during alvs , the vibrating antenniform leg is positioned close to the receiver ' s trichobothria ; alv excites these trichobothria via near - field sound and occurs at a frequency causing an especially strong and sustained response in them . as a result , alv duration can be extracted from the trichobothria response . this is the first reported match between an intraspecific near - field sound signal and the response properties of a filiform hair . we suggest that such near - field sound communication may be more common across arthropod species possessing filiform hairs than previously recognized .
male p . marginemaculatus were collected from big pine key , fl , usa , in august 2006 and housed on a reversed 12 : 12 hours light cycle . experiments were performed at 23–24°c during the subjective night .
we staged 21 agonistic encounters between eight mature male whip spiders . each animal was paired in multiple contests with previously unencountered opponents . contests were separated by more than 8 days to exclude stress or fatigue effects .
contests were staged in a 20 cm diameter circular arena with 5 cm high acetate sides . lighting was from standard fluorescent room lights and an infrared led source ( stroboled v3 , aos technologies , baden , switzerland ) to allow filming ; we noted no effects of lighting on contest behaviour . each contest was filmed at low speed from three angles ( front , back and top ) using mirrors and a horizontally mounted digital camcorder ( dcr - hc65 , sony electronics , inc . , usa ) . an alv display from each animal was also filmed simultaneously at 500fps using a fastcam 1024pci high - speed digital camera ( photron usa , san diego , ca , usa ) mounted next to the camcorder . the short high - speed recording length , and time taken to write captured video to disk , prevented us from filming alvs of both individuals in a contest .
we used low - speed films to measure contest phases observed , contest duration and alv duration . we used high - speed films to analyse the period and amplitude of antenniform leg movement during alv and the location of the display relative to the receiver ( electronic supplementary material ) .
trichobothrium recordings were made from the eight males used in behavioural experiments ( after more than four weeks ) , plus two males that matured during those experiments .
) . we recorded from both leg 3 trichobothria in separate experiments . whip spiders were fixed in a standardized standing position using plasticene and insect pins ( leg 3 tibia flexed 90° to femur and 45° to horizontal ) . recordings were made using two 50 μm copper wires insulated but for their tips ( advent research materials ltd , eynsham , uk ) , inserted through the walking leg femur . electrode wires did not interfere with trichobothria movements . recordings were differentially amplified using a standard ac amplifier and captured to disk using a power 1401 a - to - d converter and
2 v . 5 for windows ( cambridge electronic design ltd , cambridge , uk ) .
only the trichobothrium under investigation in a particular experiment was recorded : the other patella trichobothrium was removed and the tibia , tarsus and metatarsus of leg 3 were tightly covered with aluminium foil . manual stimulation of the hair , and comparison with responses to directed air puffs , identified the trichobothrium being recorded .
we used stimulus vibration frequencies of 1–120 hz . oscillation of the stimulus wire tip had a mean amplitude of 2600 μm ( measured using a high - speed camera and dissecting microscope ) . individual stimulus amplitude varied from this mean by up to 150 μm , due to flexions of the tungsten wire that we could not control . these variations did not explain overall patterns in our data ( e . g .
) , and were within the range seen during whip spider alv . the tungsten wire was positioned from anterior to posterior over the trichobothrium under investigation—30° angle to tibia , 1 mm above the trichobothrium cup and protruding 1 mm past it posteriorly—using a micromanipulator . stimuli had 5 , 20 or 90 s durations and were delivered using a 120 s interval . each trichobothrium was recorded in six whip spiders ; each received four to six presentations of each stimulus .
) simulated alv at 25 hz elicits two sizes of phase - locked action potential in recordings from pa1 . ( i ) pa1 recording , ( ii ) stimulator movement ( p , proximal ; d , distal ) . (
) the same stimulation to pa2 elicits one size of phase - locked action potential ( ap ) . ( i ) pa2 recording , ( ii ) stimulator movement . (
) adaptation of the pa1 and pa2 responses to 5 s bursts of simulated alv at varying frequency . adaptation is measured as the mean time of the last ap in a response . bars are s . e . m . and nonlinear regression lines are fitted ( pa1 small ap , grey triangles and grey line ,
) mean total ap numbers in pa1 and pa2 in response to 5 s simulated alv at various frequencies . for pa1 , both sizes of ap are considered together . lognormal curves were fitted by nonlinear regression ( pa1 , grey circles and grey line ,
. bars are s . e . m . trichobothrium data are the means of four to six stimulus replicates to each of six male whip spiders in (
whip spiders are difficult to collect and recovered post - experiment . we did not sacrifice animals to make trichobothrium measurements , but one large adult male was sacrificed for reasons unconnected with this study . we made measurements of both patella trichobothria from its removed left leg 3 using a dissecting microscope , spot flex 64mp digital camera ( diagnostic instruments , inc . , sterling heights , mi , usa ) and
discovery software ( media cybernetics , inc . , bethesda , md , usa ) .
whip spider contests staged in this study were typical of those described previously and alv was observed in 73 . 7 % ( 14 ) of contests ( electronic supplementary material ) . during alv , the displaying whip spider extended one antenniform leg towards its opponent and rapidly vibrated it from side to side in the horizontal plane (
) . alv made no audible sound and was sometimes performed simultaneously by both competitors . the contest winners performed alvs for a significantly greater fraction of total contest duration than the losers ( one - tailed
) . alvs were produced for a mean total duration of 66 . 6±13 . 1 s by the contest winners and 55 . 4±13 . 8 s by the losers . these comprised bouts of vibration from 2 to 91 s in duration ( mean = 20 . 7±2 . 1 s ) .
whip spider alv displays . ( a ) frame from a high - speed recording of two males performing alv . during alv , one antenniform leg of the signaller is extended towards the receiver and rapidly vibrated . arrows mark antenniform leg tips . ( b ) alvs are produced for a greater fraction of contest duration by contest winners than losers . for 14 contests containing alv , the fraction of contest duration that the loser performed alv was subtracted from that the winner did . box = 25th and 75th percentiles ; bisecting line = median ; whiskers = 10th and 90th percentiles ; outliers plotted . asterisks indicate data that are significantly greater ( p < 0 . 01 ) than a test mean of zero ( main text ) . ( c ) mean frequency of occurrence of different vibration frequencies in alvs from seven male whip spiders . vibration periods were measured for 30 consecutive alv cycles filmed at high speed and converted to frequency . bins are 4 hz ; bars are s . e . m . ( d ) mean alv amplitude for 30 consecutive cycles of alv recorded from seven male whip spiders in ( c ) . bins are 1 mm ; bars are s . e . m .
we successfully high - speed filmed alvs in seven whip spiders . the mean vibration cycle period was 46 . 75±6 . 46 ms , corresponding to 21 . 4 hz (
) . the antenniform leg always vibrated from side to side in the horizontal plane .
from the side , the signaller ' s antenniform leg arched downwards ; only its tip came close to the receiver ' s rearmost leg or opisthosoma ( abdomen ) . sometimes the tip of the signaller ' s antenniform leg came into close proximity with the receiver and may have contacted it ( we could not be certain of contact since the antenniform leg did not flex as during infrequent strong contact ) . the frequencies at which alv cycles might have contacted the receiver for each animal were 0 . 00 , 0 . 00 , 0 . 41 , 0 . 47 , 0 . 48 , 0 . 68 and 0 . 81 . these possible contacts normally occurred with the tibia or patella of the receiver ' s rearmost legs , or with the side of its opisthosoma , but they appeared too irregular to be the primary means of alv reception .
) . we found two trichobothria on the dorsal and the anterior patellae of the walking legs that would be particularly close to an alv display . we term these trichobothria pa1 and pa2 , respectively (
) . pa1 is on the dorsal patella , approximately 280 μm from its distal margin . in an adult male whip spider , its length was 386 μm and it was easily deflected by air currents and almost constantly in motion during observation . pa2 is more anterior than pa1 , approximately 100 μm from the patella ' s distal margin . in an adult male whip spider , its length was 193 μm and it was less easily stimulated than pa1 , requiring directed air currents . while standing , the dorsal surface of the leg is tilted posteriorly , so that the two trichobothria project rearwards ( pa1 ) and upwards ( pa2 ) .
alv occurs close to the walking leg trichobothria . ( a ) the mean proportion of alv display duration that the signaller ' s antenniform leg overlays particular receiver body regions . proportions of alv duration are indicated by : white , 0 . 00 ; light grey , 0 . 01–0 . 25 ; dark grey , 0 . 26–0 . 50 ; black , greater than 0 . 51 . the receiver is drawn from a dorsal aspect and body regions are as follows : opisthosoma ( or abdomen ; op . ) ; prosoma ( or cephalothorax ; pro . ) ; pedipalps ; proximal leg regions ( p ) ; and distal leg regions ( d ) . proportions of alv duration were calculated by recording antenniform leg position at 10 ms intervals from 1600 ms samples of alv for each of the seven whip spiders . mean proportions of alv duration and standard deviations for each body region are listed in the electronic supplementary material . data are corrected as if the signaller performed alv with its left antenniform leg . ( b ) trichobothria on the walking leg patella . patella pictured from above a whip spider in normal standing position ( leg dorsal aspect twisted posteriorly ) . f , femur ; pa , patella ; ti , tibia ; pa1 and pa2 , patella trichobothria 1 and 2 .
electrophysiological recordings showed that trichobothria pa1 and pa2 responded to direct contact ( data not shown ) . we wanted to see whether alv displays could also excite these trichobothria without contact via near - field sound . we simulated the air movements induced by alv using a device that moved a wire of the same thickness as an antenniform leg at an equivalent amplitude and frequency to real alv displays (
; electronic supplementary material ) . with this stimulus positioned as during alv , without physical contact between stimulus and trichobothrium , we recorded action potentials in pa1 and pa2 in response to simulated alv (
) . we noted two action potentials from pa1 and one from pa2 . these were phase - locked to the alv stimulus across a range of vibration frequencies ( from 1 to 120 hz ; electronic supplementary material ) . however , higher simulated alv frequencies caused faster adaptation of the responses of both trichobothria (
) . thus , the strongest response over time was induced by the highest simulated alv frequency that did not induce strong adaptation , as indicated by total action potential numbers induced in each trichobothrium by each stimulus . these counts fitted a lognormal distribution around the most common frequencies in real alvs for both pa1 and pa2 (
) , the mean time between first and last action potential in each bout was 19 . 759±0 . 068 s for the larger action potential and 19 . 934±0 . 014 s for the smaller action potential , matching the 20 s stimulus duration .
stimulation of pa1 with simulated alvs representing aspects of real displays . ( a ) peri - stimulus time histogram of pa1 response to 90 s simulated alv at 25 hz . data are plotted for both action potentials ( ap ) and are the mean of four stimulus presentations each to six male whip spiders . bins are 1 s ; bars are s . e . m . ( filled circles , small ap ; open circles , large ap ) . ( b , c ) response of pa1 mechanoreceptors in a single whip spider to repeated bouts of simulated alv with a 5 s inter - stimulus interval . bins are 1 s ( i , small ap ; ii , large ap ) .
we describe the agonistic alvs of whip spiders . these displays are produced for longer by the contest winners than the losers and have previously been proposed to convey information used to resolve agonistic contests ( fowler - finn & hebets 2006 ) . the form of an alv display is suited to providing sustained , strong excitation to the two closest trichobothria on the walking leg patellae , and thus these trichobothria can convey alv duration to the central nervous system . this is , to our knowledge , the first reported match between an intraspecific near - field sound signal and the response properties of a filiform hair .
alvs are produced by whip spiders during agonistic contests , and for a significantly greater duration by the contest winners than the losers ( alexander 1962 ; weygoldt 2000 ; fowler - finn & hebets 2006 ) . alvs occurred in 73 . 7 % of contests ; normally one individual retreated following these , and only 10 . 5 % of contests escalated to physical contact . this suggests that alv displays are signals conveying information important for resolving contest outcome .
as discussed above , the form of alv is suited to eliciting the maximum number of action potentials in the patella trichobothria over the total duration of the display . the evolutionary implications of this match are intriguing : since trichobothria have repeatedly been shown to function in predator or prey detection , and their response properties are well tuned to such a role , alv may deliberately exploit these response properties , suggesting a sensory bias origin of signal form ( e . g . basolo 1990 ; ryan 1990 ; proctor 1991 ) . the presence of trichobothria on leg regions typically stimulated during alv may even suggest that they have been co - opted to function specifically in signal detection from an original predator detection role ( e . g . conner 1999 ) .
we thank the national key deer refuge for permitting whip spider collection ; a . s . french and j . r . kelty for their help with stimulator construction ; k . d . fowler - finn , a . s . rundus , s . k . schwartz , p . s . shamble , d . j . wilgers and r . h . willemart for their comments and discussion . funding was from a searle scholar grant to e . a . h .
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. . . hence , the structure may be shaped by mate choice in the way that females use the tactical cues given by the spermatophore structure to ' assess ' the quality of the male ( peretti 9 . 7 sexual selection 148 2004 ) . in a broad comparative approach , including both insects and arachnids , it was found that spermatophore complexity generally cannot be explained with a lock - and - key mechanism , but might rather be shaped by the female choice and the demand of protection of the sperm against environmental impacts ( proctor et al . 1995 ) . though , some of the protuberances often play a role in interlocking with the female genital tract , in order to increase the duration of sperm transfer ) . . . .
. . . michalska , unpubl . ) . spermatophores can be regarded not only as a packaging for spermatozoa but also as male genitalia that fit to the genitalia of females and enable female insemination ( eberhard 1985 ; proctor et al . 1995 ; peretti 2003 ) . in eriophyoid mites , both the head and the stalk of a spermatophore appear to play important roles in the act of female self–insemination . . . .
. . . these predictions concern sympatrically living species in which reproductive character displacement is expected as a reproductive isolation mechanism against hybridization ( shapiro & porter , 1989 ) . maximum attention has been paid to the variability of genitalia between species ( eberhard , 1985 ; proctor , baker & gwynne , 1995 ) , but studies on the other three predictions are rare . several hypotheses based on sexual selection have also been proposed to explain peculiar patterns of genital evolution . . . .
. . . three main hypotheses have been proposed to explain the evolution of male genitalia : the lock - and - key , the pleiotropy , and the sexual selection hypotheses . under the lock - and - key hypothesis , selection for preinsemination reproductive isolation is predicted to favour male genitalia ( the key ) that provides an exact mechanical fit to female genitalia ( the lock ) so that male genitalia become highly canalized ( eberhard , 1985 ; proctor et al . , 1995 ; arnqvist , 1997 ) . the pleiotropy hypothesis proposes that variation in genitalic morphology is selectively neutral and that male genitalia evolve via pleiotropic effects of genes that code for both genital and general morphological characters ( mayr , 1963 ) . . . .
. . . the lock - and - key hypothesis states that genitalia evolve via selection for preinsemination reproductive isolation ( west - eberhard 1983 ; eberhard 1985 ; arnqvist & thornhill 1998 ) . the hypothesis posits that female genitalia represent a ' lock ' and male genitalia represent a ' key ' whereby the female ' lock ' allows access only to an appropriate male ' key ' , thus preventing cross - species mating ( eberhard 1985 ; proctor et al . 1995 ) . . . .
. . . the lock - and - key hypothesis states that genitalia evolve via selection for preinsemination reproductive isolation ( west - eberhard 1983 ; eberhard 1985 ; arnqvist & thornhill 1998 ) . the hypothesis posits that female genitalia represent a ' lock ' and male genitalia represent a ' key ' whereby the female ' lock ' allows access only to an appropriate male ' key ' , thus preventing cross - species mating ( eberhard 1985 ; proctor et al . 1995 ) . the findings of comparative studies ( eberhard 1985eberhard , 1992 arnqvist 1998 ) and empirical data ( goulson 1993 ; arnqvist & thornhill 1998 ) conflict with the hypothesis although it is conceivable that genital traits contribute to the initial establishment of reproductive barriers during speciation events ( shapiro & porter 1989 ; civetta & singh 1998 ) . . . .
. . . spermatophore and deposited on a substrate or onto the female ' s body ( kearn 1970 ; kearn and whittington 1992 ; spermatophores are known from a wide of barnes 1987 ; rieger et ale 1991 ; proctor et al . 1995 ) . tebrate taxa . . . .
arachnids , myriapods , and wingless hexapods exhibit a fascinating diversity of sperm transfer behaviors . modes of sperm transfer can be categorized by the degree of contact between male and female during transfer , with direct transfer ( copulation ) involving the greatest contact , paired - indirect transfer an intermediate degree , and dissociated transfer the least . internal fertilization , . . . [ show full abstract ]
water mites of the genus arrenurus vary in male sexual dimorphism and in sperm transfer behaviour . although it is a very large genus ( ≈800 spp . ) , mating behaviour has been described for only a few species . here sperm transfer behaviour is described for the first time in a north american species , arrenurus manubriator . behaviour patterns can be divided into pre - pairing ( readiness posture and . . . [ show full abstract ]
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( phd , co - supervised with r . l . baker , 1992 )
i divide sperm transfer behaviour of 23 genera of water mites ( acari : parasitengona ) into groups based on proximity of male and female during transfer , and describe spermatophore morphology of several genera ( chapter 1 ) . the great diversity of behaviour in water mites is compared to that in other arachnids , and hypotheses for this diversity are proposed . i describe functional morphology of spermatophores , and the implications of mating behaviour and spermatophore morphology for phylogenetic relationships within water mites and between this group and terrestrial acari .
in chapter 2 i develop and use a comparative test for non - reversing characters to test hypotheses about the role of habitat and swimming behaviour in the repeated evolution of copulation in water mites . evolutions of copulation among mites living in rapidly - flowing water were not more frequent than expected by chance , but there were more evolutions than expected among swimming mites . the association between copulation and swimming is not perfect , and exceptions may be due to other causes of substrate dissociation , or to males having developed alternative methods of increasing the probability of females encountering spermatophores .
( unionicolidae ) . male courtship appears to induce predatory behaviour in females . behavioural evidence includes : females responding to male trembling as they do to vibrations caused by copepod prey ; males trembling at frequencies produced by swimming copepods ; and hungry females showing more response than well - fed females to courting males . cladistic analysis of the evolution of predatory and courtship behaviour suggests that predatory behaviour arose once , and trembling either evolved simultaneously or twice thereafter .
the female - choice hypothesis for spermatophore complexity predicts that spermatophores from species with close male / female contact during sperm transfer should be more complex than those without such contact . in chapter 4 i test this using volunteers unaware of the hypothesis to rank the complexity of spermatophores from a phylogenetically - controlled group of species , including several water mites . results do not support female choice as a general explanation for spermatophore complexity .
the following articles are merged in scholar . their combined citations are counted only for the first article .
this
cited by
count includes citations to the following articles in scholar . the ones marked
cp catterall , j kanowski , gw wardell - johnson , h proctor , t reis , . . .
i galván , e barba , r piculo , jl cantó , v cortés , js monrós , f atiénzar , . . .
this revised version was published online in july 2006 with corrections to the cover date .
baker g . t . 1996 . chemoreception in four species of water mites ( acari : hydrachnida ) : behavioural and morphological evidence . exp . appl . acarol . 20 : 445 - 455 .
( thon ) , 1899 ( hydrachnellae , acari ) . zool . jahrb . system . 89 : 501 - 584 .
: males capitalize on female adaptations for predation . anim . behav . 42 : 589 - 598 .
proctor h . c . 1992a . mating and spermatophore morphology of water mites ( acari : parasitengona ) . zool . j . linn . soc - lond . 106 : 341 - 384 .
proctor h . c . 1992b . sensory exploitation and the evolution of male mating behavior : a cladistic test using water mites ( acari : parasitengona ) . anim . behav . 44 : 745 - 752 .
proctor h . c . and smith b . p . 1994 . mating behaviour of the water mite
marshall ( acari : arrenuridae ) . j . zool - lond . 232 : 473 - 483 .
proctor h . c . and wilkinson k . 2001 . coercion and deceit : water mites ( acari : hydracarina ) and the study of intersexual conflict . in : halliday r . b . , walter d . e . , proctor h . c . , norton r . a . and colloff m . j . ( eds ) ,
smith b . p . 1998 . loss of larval parasitism in parasitengonine mites . exp . appl . acarol . 22 : 187 - 199 .
sonenshine d . 1985 . pheromones and other semiochemicals of the acari . ann . rev . entomol . 30 : 1 - 28 .
witte h . 1984 . the evolution of the mechanisms of reproduction in the parasitengonae ( acarina : prostigmata ) . in : griffiths d . a . and bowman c . e . ( eds ) ,
. vol . 1 . ellis horwood ltd . , chichester , uk , pp . 470 - 478 .
witte h . 1991 . indirect sperm transfer in prostigmatic mites from a phylogenetic viewpoint . in : schuster r . and murphy p . w . ( eds ) ,
( johnston ) 1852 ( acari : prostigmata : parasitengonae ) . acarologia 37 : 201 - 209 .
smith , b . p . & hagman , j . exp appl acarol ( 2002 ) 27 : 257 . urltoken
artificial life models have consistently implemented communication as an exchange of signals over dedicated and functionally isolated channels . i argue that such a feature prevents models from providing a satisfactory account of the origins of communication and present a model in which there are no dedicated channels . agents controlled by neural networks and equipped with proximity sensors and wheels are presented with a co - ordinated movement task . it is observed that functional , but non - communicative , behaviours which evolve in the early stages of the simulation both make possible , and form the basis of , the communicative behaviour which subsequently evolves .
these keywords were added by machine and not by the authors . this process is experimental and the keywords may be updated as the learning algorithm improves .
d . ackley and m . littman . altruism in the evolution of cooperation . in r . brooks and p . maes , editors ,
h . kawamura , m . yamamoto , k . suzuki , and a . ochuchi . ants war with evolutive pheromone style communication . in d . floreano , j - d . nicoud , and f . mondada , editors ,
m . levin . the evolution of understanding : a genetic algorithm model of the evolution of communication .
b . maclennan . synthetic ecology : an approach to the study of communication . in c . langton et . al . , editor ,
m . quinn . a comparison of approaches to the evolution of homogeneous multi - robot teams . in
c reynolds . flocks , herds , and schools : a distributed behavioral model .
g . werner and d . dyer . the evolution of communication in artificial organisms . in c . langton et . al . , editor ,
quinn m . ( 2001 ) evolving communication without dedicated communication channels . in : kelemen j . , sosík p . ( eds ) advances in artificial life . ecal 2001 . lecture notes in computer science , vol 2159 . springer , berlin , heidelberg
morphological ( or other phenotypic ) differences between conspecific males and females in characters other than their directly reproductive structures .
• do not directly y * participate in the sexual - reproductive process . • are usually involved in mating opportunity and / or mating success .
throat patch is inflated as part of male mating display , females lack red throat patch .
pollen - producing ( male / staminate ) flower larger and / or more attractive to pollinators than ovary producing ( female / carpellate ) flower .
this form of selection depends , not on a struggle for existence in relation to other organic beings or to external conditions , but on a struggle between the individuals of one sex , generally the males , for the possession of the other sex .
• intrasexual competition or mate - choice . - - enhance reproductive success of individuals bearing the characteristics . • many sexually selected characters do not t * appear to be adaptive in the general sense . • may increase mating success and d * be otherwise adaptive .
increase risk ( of predation ) or energetic cost . i . e . conspicuous coloration / plumage , fighting for females , large ornamentation
evolutionary
trade offs
the advantage gained in reproductive success must be greater than the reduction in survival .
1 . the relative parental investment ( the
cost
of reproduction ) 2 . the reproductive limiting resource ( frequently different for females vs . males )
neither sex - no
cost
of parental care . ( birds and mammals are the most obvious exceptions )
• parental care is an added parental investment • is an additional consideration in the
cost
of reproduction • can significantly alter or even reverse the pattern of sexual selection on males vs . females
• fitness of their mates ( limiting reproductive resource ) - is reduced by mating with
poor quality
males . - female ' s mate - choice will affect her fitness .
females - high parental investment males - low parental investment ( neither parent cares for the young . )
slope of the best - fit line for the covariance of fitness ( here , number of offspring ) versus a sexual character ( here , number of mates ) .
selection gradient for sexual selection • covariance of fitness vs . a sexual character • slope best fit line estimates strength of sexual selection
one male / one female exclusive mating pairs . - sex ratios tend to be 1 : 1 - little competition for access to females - sexes tend to exhibit little sexual dimorphism
• intense competition among males for access to females . • sexual selection will favor male characters that contribute to greater access to females . • sexual dimorphism is typically more strongly developed in more polygynous species .
in species in which one sex directly control access to females or to critical resources .
• competition among members of the same sex . • determines who mates . •
winners
mate more often • sexual selection is for traits associated with competition among members of the sex subject to strong sexual selection .
- males
win
mates by advertisement or other indirect competition - sexual selection is for traits associated with
attracting
females - females are
choosy
: mate with the
most attractive
males ( assortative mating ) - the selection is between the members of the two sexes ( intersexual ) .
- sexually dimorphic for color and tail length . - long tails are physiologically costly . - females prefer ( choose ) males with longer tails .
1 . advertisement of higher fitness 2 . female acquisition of resources 3 . female choice based on preexisting sensory bias 4 . selection reinforcing arbitrary female preferences
• sexual selection for traits that are correlated with higher fitness of the male - can include many different types of traits examples : morphological , pheromonal , vocal , behavioral ( i . e . , courtship behaviors )
- variation in male in vocalizations - females
prefer
males with longer , more rapidly repeated calls - experiments show significantly higher
fitness
in the offspring of males with longer , more rapidly repeated calls .
• female choice may be a by - product of a behavior associated with some other aspect of survival ( i . e . , predator avoidance , food seeking , identification of conspecifics ) - female preference evolves first , corresponding male character is reinforced by the female preferences
offspring that receive genes for the trait value ( from their fathers ) also receive the genes for the preference of that trait value ( from their mother ) . sons of mothers with a preference for a male trait value (
sexy sons
) have sisters who inherit the preference for that trait value .
males with that trait value (
sexy sons
of choosy mothers ) will be chosen for reproduction more often than males of the optimal - survival trait value .
parental investment for the male may be equal to or even greater than that for the female .
- access to mates will be a limiting resource for females . - females will compete for access to males and exhibit showy dimorphisms . - males will exhibit mate choice .


the telefomin cuscus ( phalanger matanim ) is a probably extinct possum found on new guinea .

it is named after the telefol ethnic group , who knew the animal long before it was discovered scientifically by the australian zoologist tim flannery .

it lives in the oak forests along the nong river in the middle of papua new guinea , at altitudes between 1500 and 2000 m .

after the drought and wildfires of 1997 , these forests were destroyed , which probably made the species extinct .

telefomin cuscus

the same cannot however be said for some of its relatives . several species of phalanger are widespread and share the ground cuscus’ status , but several species are confine to islands or seem to be geographically isolated , and these are in far more trouble . the telefomin cuscus
two species in this family are considered endangered and at risk of going extinct in the wild . these are the black - spotted cuscus and the telefomin cuscus . the population of black - spotted cuscuses is declining because of habitat loss and continued hunting . little is known about the telefomin cuscus . two other species are considered vulnerable , facing a high risk of extinction , and half a dozen others are of concern to conservationists , but too little is known about them to make an accurate population evaluation .
geographic range : this cuscus is found in new guinea and the aru islands .
physical characteristics : the ground cuscus has fur that is light to dark gray with a stripe down its back .
most members of this family are nocturnal , or active at night , but the black - spotted cuscus and the sulawesi bear cuscus feed during the day . all species are arboreal ( tree - dwelling ) except for the ground cuscus—but even though this animal lives in burrows underground , it is a good climber , and climbs trees to feed on fruit .
conservation status : the ground cuscus is common in many parts of new guinea , and is probably not threatened . ∎
cuscuses are hunted for meat and sometimes fur in new guinea . some species , such as the common spotted cuscus , are also sold as pets . cuscuses play a role in religious beliefs in some parts of indonesia , and in these areas , they are not eaten . the common brush - tailed possum is considered a pest in many areas . the telefomin cuscus was not discovered until the late 1980s , and so it is of special interest to scientists .
members of this family have soft , dense fur that hides small ears . most species are a solid brown or gray , but the woodlark cuscus and the black spotted cuscus are spectacularly patterned . many other species have a dark stripe that runs down the top of their back .
other critically endangered mammals in indonesia include sir david’s long - beaked echidna , thr eastern long - beaked echidna , the western long - beaked echidna , the talaud bear cuscus , the telefomin cuscus , the black - spotted cuscus , the blue - eyed spotted cuscus , the northern glider , the golden - mantled tree - kangaroo , the tenkile , the black dorcopsis , the sumatran water shrew , jenkin’s shrew , the flores shrew , bulmer’s fruit bat , the aru flying fox , the black - eared flying fox , the sunda pangolin , the sumatran rhinoceros , the heavenly hill rat , the manusela mosaic - tailed rat , the lowland brush mouse , the enggano rat , the biak giant rat , emma’s giant rat , the celebes crested macaque , the pagai island macaque , the sarawak surili , and the pig - tailed langur .
diet : the ground cuscus eats mostly fruit , but will also eat leaves and sometimes insects and small vertebrates ( animals with a backbone ) .
has a similar status to the ground cuscus , and like it has several close relatives with restricted ranges that are in a more serious state .
the phalanger family phalangeridae is one of the few groups of australian mammals to successfully cross wallaces line that separates australasian animals from those of mainland asia . the bear cuscus
possums and cuscuses range in size from 24 to 47 inches ( 60 to 120 centimeters ) long , including the tail , and weigh from 2 to 22 pounds ( 1 to 10 kilograms ) . the smallest member of this family is the small sulawesi cuscus , and the largest is the sulawesi bear cuscus . both live in indonesia .
behavior and reproduction : this is the only cuscus that sleeps in underground burrows and moves along the rainforest floor . its burrows are usually under trees , along streams , or in caves . the ground cuscus is active at night and searches for food on the ground and in trees . the young leave their mother ' s pouch five to seven months after birth .
is only known from six individuals and the only site where it was known to occur was completely destroyed by fire in 1998 , and it is either already extinct or critically endangered . the woodlark cuscus
( also 2 species ) , which occur on the same islands , although not necessarily in the same habitat . with a range from sulawesi through new guinea into northern australia , the common spotted cuscus
this species is known from the areas of telefomin and tifalmin , papua new guinea . the species might be found further north - east or west of the known collection localities . the only place where it is known with certainty was largely destroyed by fire in 1998 . it has been recorded from between 1 , 400 and 2 , 600 m asl ( 1 , 400 - 1 , 800 is the core elevation range ) . it occurs sympatrically with the other species .
which is widely spread throughout australia , and as an introduced species is a major threat to new zealand’s native fauna , the ground cuscus is currently listed as least concern , although deforestation and over - hunting may be a cause of local declines . it is quite a widespread species however , so as a species it is not in any especial trouble .
as a fairly unspecialised marsupial , the ground cuscus has a pouch where it retains its single baby for around 3 months , after which it takes another 2 – 4 months to become independent , at which point it leaves to join the adult population . they breed all year round , and the actual pregnancy only takes 13 days . lifespan in the wild is unclear , but in captivity they can live to be 15 years old .
the common spotted cuscus was selected to be one of twelve endangered species featured on a 2001 united nations 34 cent stamp . every year since 1993 , the united nations has released a new series of stamps in an effort to bring attention to endangered species and to cites ( the convention on international trade in endangered species ) , an agreement among nations to help preserve species by controlling their exportation and importation ( http : / / www . cites . org ) .
all the phalangerids seem to have a fairly similar diet , being basically generalists feeding on flowers , fruits , nectar and a varying amount of animal food . most are about the size of a domestic cat , and tend to live mainly in trees – the ground cuscus is unusual in using terrestrial burrows , although even it spends a lot of time in trees . in some ways cuscuses can be thought of as the ecological equivalents of primates , although unlike them ( which are of course placental mammals ) they are solitary and nocturnal by nature and are not renowned for their intelligence .
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justification : listed as critically endangered ( possibly extinct ) because its only known location was completely destroyed by fire in 1998 . there is a strong possibility that this species is extinct , however , surveys of other potential sites are required . should this species be located , its total population would be very small ( probably less than 50 mature individuals ) .
it is known from only six specimens . an individual that is presumed to be this species was captured in 1997 ( l . salas pers . comm . 2015 ) , but was not collected . the area in which the species occurs has not been well surveyed . if the species still exists , it is likely that its population is 50 or fewer mature individuals .
it is found in mid - montane tropical moist oak forest and has a narrow altitudinal range . little more is known about the natural history of this distinctive species .
the habitat at the type locality was completely burnt in the 1998 el niño event . this species is threatened by local hunting for food and loss of suitable habitat through human encroachment .
it is not known if the species is present in any protected areas . further field surveys are needed to confirm the existence of this species . hunting regulations may be needed to prevent its overexploitation .
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english - german online dictionary developed to help you share your knowledge with others . more information ! contains translations by tu chemnitz and mr honey ' s business dictionary ( german - english ) . thanks on that account ! links to this dictionary or to single translations are very welcome ! questions and answers
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wilson , d and reeder , d . ( eds . ) 2005 . mammal species of the world . a taxonomic and geographic reference . 3rd ed . - - 2142 pp . johns hopkins university press .
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a megadiverse country facing extreme environmental challenges and habitat losses , indonesia has many critically endangered mammals calling it home .
indonesia is a megadiverse country facing extreme environmental challenges and habitat loss , an unfortunate effect extending to its many critically endangered mammals . indonesia’s numerous islands are home to some of the world’s iconic and endemic mammals . some of these mammals are however endangered from an environmental threat such as habitat loss and poaching .
only inhibits the island of java in indonesia . the leopard’s color can either be entirely black or orange with spots . the leopard has a wide habitat range on the island , from dense tropical rainforest to dry deciduous forests . the javan leopard feeds on other animals such as wild boar , deer , javan gibbon , silvery lutung , and long - tailed macaques . the leopard also preys on domestic animals such as poultry and dogs . the javan leopard is one of the world’s most threatened big cat species . the leopard’s sustainability is continuously threatened by poaching , human encroachment , extensive agriculture and loss of habitat . population pressure has caused 90 % of java’s natural vegetation to be cleared . the leopard’s numbers is estimated at less than 250 mature individuals and is already cause for environmental alarm . the leopard is conserved in several national parks such as ujung kulon national park , merapi national park , gunung halimun national park , and baluran national park . poaching of the animal has subsequently been made illegal .
the sumatran tiger ( panthera tigris sumatrae ) is the smallest tiger subspecies , and is only found on the indonesian island of sumatra . the tiger has an orange coat with black stripes . the tiger’s habitat ranges from lowland forest , mountain forest to tropical rainforest , and swamp forests . the tiger has slightly webbed paws , which helps it to swim in pursuit of prey . its diet mainly consists of wild boar , deer , tapir , monkeys , cattle , and fish . the leopard hunts mostly at night . the tiger has been on the international union for the conservation of nature ' s ( iucn ' s ) red list as a critically endangered species since 2008 . the sumatran tiger is continuously threatened by habitat loss and hunting . the tiger is conserved in five national parks and two game reserves , the largest numbers being in the gunung leuser national park . about 100 are believed to be unprotected in the world , making them susceptible to human interference .
the javan rhinoceros ( rhinoceros sondaicus ) is one of the most threatened rhinos on earth , and one which is found on the island of java in indonesia . the animal’s skin is scale - like and ranges from gray - brown to gray , and it has one gray or brownish color horn . the rhinoceros has a long pointed upper lip to assist in food grasping , and it is herbivorous . the javan rhinoceros is estimated to live for an average of between 35 to 40 years while in the wild . the rhinoceros has a great sense of smell but it is short - sighted , and it is , for the most part , a solitary animal . the rhinoceros had a wide habitat range , both in the lowland and highlands and it currently exists as a single population in ujung kulon peninsula , on java island . the largest threat to the rhinoceros has been poaching for their horns coupled with habitat degradation . the ujung kulon peninsula remains protected to sustain their already dismal numbers . poaching of the rhinoceros is also illegal on java island .
the sumatran orangutan ( pongo abelii ) is a species of the orangutan primates inhabiting the indonesian island of sumatra . there , the species is mainly concentrated on the highly significant leuser ecosystem . recent statistics estimate the species’ population to be at 14 , 613 . the animal is characterized by pale red long hair and a long face . the sumatran orangutan feeds on insects and fruits as well as bird eggs . the species makes tools from branches for use while searching for food . the animal spends most of its time on trees because of predators and has adopted various locomotion techniques due to its heavy size . the animal has five life cycles and can live for more than 50 years . the species is listed as critically endangered and is threatened by continued degradation of the leuser ecosystem . logging , human encroachment , and mining activities have adversely affected the species’ population . sumatran orangutan inhabits the gunung leuser national park and the bukit lawang , where conservation programs have been established .
conservationists issued a chilling warning this week telling new scientists magazine that global wildlife populations are set to have fallen by more than two thirds of 1970 levels by the year 2020 – and there is no sign that declining wildlife numbers will slow down in our lifetime .
researchers say mammal , bird , fish , amphibian , and reptile populations have fallen by 58 % from 1970 to 2012 . at the current rate ( about 2 % reduction per year ) , by 2020 , the populations will have declined by more than 67 % . this has prompted experts to warn that we are facing a global “
“for the first time since the demise of the dinosaurs 65 million years ago , we face a global mass extinction of wildlife . we ignore the decline of other species at our peril – for they are the
that reveals our impact on the world that sustains us . humanity’s misuse of natural resources is threatening habitats , pushing irreplaceable species to the brink and threatening the stability of our climate . ”
“we know how to stop this . it requires governments , businesses , and citizens to rethink how we produce , consume , measure success and value the natural environment . ”
wildlife populations are impacted by poaching , disease , over - hunting , and manmade changes to the animals’ habitat from mining , climate change , pollution , and other human activities . below is a list of some of the animals that have recently gone extinct thanks to careless human activities .
last existed in cameroon . extinct due to poaching ( there was a high demand for the rhino’s horn ) .
went extinct in 2000 due to over - hunting . scientists tried to clone a pyrenean ibex in 2009 but it died from lung defects shortly after birth .
once represented 25 - 40 % of the bird population in the united states . habitat was destroyed as humans cleared forests for farmland . with nowhere to go , passenger pigeons decimated farmers’ crops . angry farmers began shooting the birds for food . the last passenger pigeon ( named “martha” ) died at the cincinnati zoo in 1914 .
a subspecies of the plains zebra , it was hunted for its unusual hide . the last known quagga died at the amsterdam zoo .
these cute little guys were hunted to extinction , exploited for their fur , meat , and oil by fishermen and whalers . coastal development in the caribbean sea and gulf of mexico destroyed their habitat . they were declared extinct in 2008 .
was once found along the coasts of maine and new brunswick . prized for its fur , it was carelessly hunted to extinction .
once lived along the australian continent but as their habitat was destroyed by humans , they migrated to the island of tasmania . they were often shot and trapped by the locals . they were last seen in the 1940’s .
oddly , the tecopa pupfish lived in the mojave desert in california , surviving in waters as warm as 108 degrees . human development left their habitat unsurvivable and they were declared extinct in 1970 .
a beautiful flightless coastal bird that lived around the north atlantic ( canada , greenland , iceland , british isles , and scandinavia ) . they were slaughtered for their feathers and driven to extinction .
also known as the bubal antelope , they lived in north africa . they were hunted to extinction sometime around 1950 .
declared extinct in august 2007 . the golden toad was driven to extinction by pollution and
the last known spix macaw was seen in 2000 . their extinction is attributed to hunting , destruction of habitat , and the introduction of africanized bees ( aka killer bees ) into their habitat ( the bees raided their nests ) .
declared extinct in 2008 . the reason for extinction is believed to be loss of habitat .
last seen in 2004 , is believed to be extinct due to habitat destruction and global warming .
valued for its skin ( used to make gloves and handbags ) , it was last seen in 2002 and believed to be extinct due to overfishing , boat traffic , habitat loss , and pollution .
native to costa rica , it was declared extinct in 2004 . its extinction is likely due to amphibian disease and the effects of climate change .
declared extinct in 2010 due to habitat destruction . only one photograph exists of the beautiful bird .
the last pinta island tortoise , named lonesome george , died in a research pen in 2012 . his age was believed to be over 100 years old . the species was driven to extinction due to loss of habitat .
once living on the canary islands off the coast of west africa , they were declared extinct in 1994 . only 4 specimens of stuffed birds exist in museums today .
after loss of habitat ( swamps ) in the southern united states , the last definitive sighting was in 1943 .
habitat loss from draining of marshes for farming and homes , the last pair of ducks were spotted in the wild in 1979 . the last known pair died in captivity at sea world in san diego in 1981 .
once found all over the east coast of florida , they were declared extinct in 1990 . the species died out from ddt pesticide spraying in its habitat .
the last of this species died in a laboratory in 1983 . unusually , the frog swallowed its eggs and birthed them out of their mouth . the mother had the unique ability to shut off her stomach
while carrying the young in her stomach . scientists had hoped to learn something about how to cure human ulcers from the frog before it vanished from our planet .
nearly 20 , 000 species of plants and animals are at a high risk of extinction . mammals have faired far better than other classes such as birds and fish . as an example illustrating the seriousness of the problem , below is a complete list of recent mammal extinctions . other lists of extinct animal classes ( e . g . birds , amphibians , fish ) are much longer .
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the phalangeridae family , or phalangers ( fah - lan - jerz ) , are made up of five groups of species . three of these groups are cuscuses and two are possums . phalangers are small - to medium - sized marsupial mammals . like all marsupial mammals , the females give birth to tiny , underdeveloped young that finish their development in their mother ' s pouch .
all cuscuses and possums are good climbers . their feet are adapted to life in the trees . their hind feet have five toes . the first toe ( called the hallux , hal - lux ) has no claw , and is opposed to the other four . this means that this toe , like the thumb on a human hand , can reach across and touch the tip of the other toes ( unlike , for example , a dog paw or human foot , where none of the toes can bend to touch each other ) . the first two toes on the front feet are also opposable . this adaptation makes it easier to grip branches when climbing . possums and cuscuses also have a prehensile , or flexible grasping , tail that they can wrap around branches to help steady themselves . usually the tail has no fur on it to improve its grip .
phalangers are found in new guinea , australia , tasmania , the indonesian island of sulawesi , and a few other small islands . the common brush - tailed possum was introduced in new zealand over a century ago and has become an alien ( non - native ) pest species .
possums and cuscuses spend their lives in trees . most live in rainforests . however , the common brush - tailed possum has adapted to life in developed areas . it is often found in suburban gardens and city parks . sometimes it becomes a pest when it makes its home in buildings by finding openings in the roofline and nesting between the house ceiling and the roof .
possums and cuscuses are herbivores , eating almost exclusively plants . some eat mainly leaves , while others eat mainly fruit . the common brush - tailed possum eats a wider variety of foods than most members of this family , adapting its diet to what is abundant in any given area .
little is known about the social behavior of members of this family . most species appear to live alone , although a few may form pairs . males are aggressive toward each other when their home range overlaps . females usually produce two litters consisting of one offspring each year . like all marsupials , the young are tiny , undeveloped creatures that finish maturing while attached to a teat , or nipple , in the mother ' s forward - facing pouch . after five to eight months , the young leave the pouch and are carried on their mother ' s back for a few more weeks or months .
habitat : this animal lives in the rainforest from sea level to 8 , 900 feet ( 2 , 700 meters ) .
ground cuscuses and people : these cuscuses are hunted throughout new guinea , and are important figures in local folklore in some areas .
physical characteristics : the common brush - tailed possum varies widely in size and color . its fur can be black , gray , reddish , or brown . in the colder parts of its range , individuals tend to be larger and furrier than those who live in warmer regions . unlike some members of this family , the common brush - tailed has a patch of bushy fur on its tail .
geographic range : brush - tailed possums live in eastern and southwestern australia and in new zealand .
habitat : the brush - tailed possum is adaptable , living in cool , damp forests and dry regions with few trees . it has adjusted successfully to life in city parks and the suburbs .
diet : common brush - tailed possums are herbivores , eating leaves , buds , flowers , and fruits , garden plants , herbs , and grasses .
behavior and reproduction : common brush - tailed possums are active at night and normally live alone . however , if there are many possums and few places to shelter , they may share their sleeping space with another possum . male common brush - tailed possums try to avoid conflict with other males , although they can be aggressive in defending their home range . they are known for their loud grunts , growls , and screeches that are used to warn away other males during breeding season . females usually have one offspring each year , born after an eighteen - day pregnancy . the young then live in the mother ' s pouch for about seven months .
common brush - tailed possums and people : this animal probably has more contact with people than any other australian marsupial because it has adapted so well to cities and suburban areas . it is trapped for its fur and is considered a pest in some farming areas and in new zealand , where it was introduced about 150 years ago .
conservation status : the common brush - tailed possum is common within its range and is not threatened with extinction . ∎
finney , tim f . mammals of new guinea , 2nd ed . ithaca , ny : cornell university press , 1995 .
menkhorst , frank . a field guide to the mammals of australia , 2nd ed . oxford , u . k . : oxford university press , 2001 .
nowak , ronald . m . walker ' s mammals of the world . baltimore : johns hopkins university press , 1995 .
parks and wildlife service tasmania .
brushtail possum .
urltoken ( accessed on june 30 , 2004 ) .
queensland government environmental protection agency / queensland parks and wildlife service .
nature conservation .
urltoken ( accessed on june 30 , 2004 ) .
possums and cuscuses : phalangeridae .
grzimek ' s student animal life resource . . encyclopedia . com . ( july 9 , 2018 ) . urltoken
possums and cuscuses : phalangeridae .
grzimek ' s student animal life resource . . retrieved july 09 , 2018 from urltoken urltoken
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( 2 species ) is known from sulawesi and nearby islands , and very little is known about it . even less known are its smaller relatives the dwarf cuscuses
as you would guess from the blog title , i am a volunteer at a zoo , where i have been volunteering for many years . in real life i am an accountant
* indarctos * a great many predators lived in europe in the glorious warmth of the mid miocene , when the continent was lush with subtropical vegetation and t . . .
let’s be honest : anoles are fascinating ! these charismatic and well - adapted lizards are always a pleasure to watch and document . better yet , no matter how . . .
in an earlier post , i introduced you all to the fusulinids , a group of complex foraminiferans that were abundant during the later palaeozoic . in that post , . . .
this article will soon be available in spanish inspired by mike van valen ' s
the ratsnake mess for dummies
please note that the information in this article . . .