pulvinitidae is a family of saltwater clams , marine bivalve molluscs in the order pterioida .

these bivalves are related to the scallops and oysters .

originally believed to be extinct and known only from fossil records , non-fossil shells of members of this family were first discovered in 1913 by the australian research vessel hms endeavour off the coast of victoria .

sixty years later , live specimens were finally discovered on the wing of a wrecked airplane at a depth of over 400 meters .

pulvinitidae

animals belonging to the bivalve family pulvinitidae were originally known only from fossil records dating back millions of years ago .
palmer , t . j . 1984 . revision of the bivalve family pulvinitidae stephenson , 1941 . palaeontology , 27 , 4 , 815–824 .
in 1984 , t palmer revised the pulvinitidae family after the new discoveries and moved the dead specimens from the ' endeavor ' to the fossil genus pulvinites .
four complete specimens of a bivalve belonging to the genus pulvinites blainville , 1824 ( pterioidea , pulvinitidae ) have been collected in the danian ( lower paleocene ) reefal linestone of vigny ( val - d’oise , france ) . pulvinites ( s . s . ) . . . more
four complete specimens of a bivalve belonging to the genus pulvinites blainville , 1824 ( pterioidea , pulvinitidae ) have been collected in the danian ( lower paleocene ) reefal linestone of vigny ( val - d’oise , france ) . pulvinites ( s . s . ) dysporista n . sp . is described . this is the first report of the member of family pulvinitidae in the european cenozoic , the only other post - mesozoic species records of pulvinites being from danian ( lower paleocene ) of egypt , from thanetian ( upper paleocene ) of california and virginia as also in recent of australia . its specific content and stratigraphical range are also briefly analyzed .
reconsideration of the characteristics of all species referred to the three genera which have hitherto constituted the family pulvinitidae stephenson , 1941 , indicates that all should be accommodated in the nominate genus , pulvinites blainville , 1824 . a new species , p . mackerrowi from the middle jurassic of england ( the earliest known representative of the family ) is described .
note : the review of the recent pulvinitidae by boss ( 1982 : 1115 ) undoubtedly refers to pulvinites exempla , but is not included in the synonymy because it does not contain the species name . likewise , franc ( 1960 : 2081 ) did not explicitly name the species in his treatment of foramelina that he placed in the family isognomonidae woodring , 1925 .
no further specimens were found for over 60 years . then in 1980 , a couple of pulvinitidae shells were brought up from a depth of 400 - 460 m off the coast of sydney by commercial trawlers . this discovery was closely followed in july 1981 by a jackpot find - a whole aeroplane wing with living specimens attached , trawled from a depth of 406m also off the coast of sydney .
pulvinites exempla , the only living species in its family and genus , represents our entire concept of the bivalve family pulvinitidae . this detailed analysis of the anatomy of p . exempla has considerably expanded our knowledge of the enigmatic bivalve family pulvinitidae and provides important clues for unveiling the evolutionary history of this ancient pterioidean lineage . many aspects of the anatomy and shell morphology prove to be unique among recent pterioidea and , more broadly , living bivalves . no specimens of p . exempla have ever been collected alive and future research is needed to complement the present work with ecological studies and observations on living animals . knowledge of the biology and morphology of this ‘living fossil’ is unquestionably critical for inferring the evolution of the family , as well as pterioidea as a whole , in light of ever - increasing palaeontological data .
fortunately in this case , the skipper with the ripped nets did not immediately throw the offending junk back overboard , but brought it back into port and contacted a shell collector about the animals living on the wing . when the collector realised that he did not recognise the bivalves , he contacted the australian museum . as a result , the first known living specimens of the pulvinitidae family made their way to the australian museum in good condition .
the microstructure of the shell is prismatonacreous with a thin outer layer and considerably thicker inner layers . the outer shell layer ( fig . 7 ) is composed of calcitic , vertical , polygonal , regular simple prisms , each approximately 12–15 µm in width . the prisms are predominantly pentagonal in cross - section , but vary greatly in both shape and size , sometimes having curved sides . individual prisms are also somewhat irregular , varying in width across height . the mineralogy is assumed to be calcitic , as shown for fossil pulvinitidae and the pterioidea in general ( carter , 1990 ) . the projecting lamellae of the outer shell surface are produced by branching of this layer ( fig . 7g ) . irregular splitting of the prismatic layer occurs parallel to the shell surface , resulting in the formation of small , empty , internal pockets ( fig . 7h ) . the prismatic layer is approximately 63 µm , whereas the nacreous layer is approximately 360 µm wide , as seen in the cross - section made at approximately one - third valve height from the ventral margin . a thin layer of periostracum covers the exterior of the prismatic layer ( fig . 7b , d , e , f ) .
the ligament is multivincular , amphidetic , and submarginal . ( despite the fact that the larger portion of the ligament is above the hinge axis , the working part of the ligament is exclusively submarginal . ) the placement of the hinge plate relative to the umbo and the deposition of resilia anterior and posterior to the umbo ( see below ) necessitate a change in the diagnoses of the pulvinitidae of palmer ( 1984 ) and ward & waller ( 1988 ) characterizing the hinge as anterior to the umbo and ligament as prosodetic . the fibrous components of the ligament are deposited in resilifers , slightly ventrally widening subtriangular ( in side view ) narrow grooves , which are rounded in cross - section . the resilifers are monoserial and breached along the ventral surface ( fig . 10b ) . no abandoned resilifers were observed . the resilifers are slightly curved towards the umbo in their dorsal part , but run nearly in parallel along most of their length . they differ in height , the tallest one being immediately ventral to the umbo , whereas the neighbouring resilifers gradually diminish in height anteriorly and posteriorly following the slopes of the hinge plate ( fig . 10a ) . the number of resilifers reaches 27 in the largest adult ( ams c . 170923 ; figs 1c , d ) . the area of the hinge plate bordering the resilifers is flat and covered by the lamellar component of the ligament , resulting in a continuous alternating pattern of lamellar and fibrous layers ( fig . 10a ) . the fibrous ligament ( the resilium ) extends slightly more ventrally than the lamellar ligament . both fibrous and lamellar layers are continuous across valves .
so it was somewhat of a surprise in 1913 when some pulvinitid specimens were trawled from a depth of 180 - 460 m off the eastern coast of victoria , by the commonwealth fisheries research vessel ' endeavour ' . although only dead shells , they were obviously of recent origin and were named foramelina exempla by the australian museum curator at the time , charles hedley .
the first question was easy - after world war ii , surplus military equipment was dumped in deep waters out to sea beyond the reach of the commercial fishing of the time . within 35 years however , with the improvement of fishing technology , australian trawlers were regularly working at these depths . trawlers began to come across the dumped equipment often tearing their trawl nets on it .
the next two questions took longer to answer . in 1997 , underwater video showed live pulvinitids attached to steep rock walls at a depth of 150 - 285 m in fiordland , new zealand . trawlers generally avoid steep rocky areas , explaining why virtually no specimens had appeared in commercial or research trawls . the plane wing was just in the right place at the right time providing a suitably hard surface for the animals to attach to .
confused by a class within a class or an order within an order ? please see our brief essay .
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 .
disclaimer : the animal diversity web is an educational resource written largely by and for college students . adw doesn ' t cover all species in the world , nor does it include all the latest scientific information about organisms we describe . though we edit our accounts for accuracy , we cannot guarantee all information in those accounts . while adw staff and contributors provide references to books and websites that we believe are reputable , we cannot necessarily endorse the contents of references beyond our control .
this material is based upon work supported by the national science foundation grants drl 0089283 , drl 0628151 , due 0633095 , drl 0918590 , and due 1122742 . additional support has come from the marisla foundation , um college of literature , science , and the arts , museum of zoology , and information and technology services .
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parent taxon : malleidae according to j . g . carter et al . 2011
see also bieler and mikkelsen 2006 , nevesskaja 2009 , spencer et al . 2004 and vokes 1980
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the symbols k . a . c . f . m . an . are used to indicate the geographical range of the species . they have been adopted to give an approxomation of the range of each species within new zealand .
( lima ( limatula ) hodgsoni e . a . smith ) ( = squamilima fleming )
( broderip in broderip & g . b . sowerby , 1835 ) ( =
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hedley had also pointed out that p . exempla resembles species of the mesozoic genera hypotrema orbigny , 1853 and pulvinites blainville , 1824 much more closely than any living bivalves because they too possess both the multivincular ligament and the foramen . however , he disregarded the phylogenetic affinity of the recent and fossil taxa based on an erroneous assumption of orbigny ( 1853 ) that the foramen in hypotrema accommodated the passage of a muscle rather than a byssus ( hedley , 1914 ) .
nearly 70 years after the endeavour expedition , another remarkable discovery was made . in july 1981 , an entire aeroplane wing ( part of military surplus dumped into the ocean after world war ii ) was trawled from a depth of 406 m off the coast of sydney with a thriving population of living foramelina exempla ( see loch , 2004 ) . however , the importance of the find escaped the attention of subsequent bivalve investigators and the shells together with the soft tissue remained unacknowledged at the australian museum in sydney ( ams c . 129659 ) .
despite a considerable interest in the group on the part of palaeontologists and neontologists , most aspects of pulvinitid shell and ligament morphology remain fragmentary , and the soft anatomy is entirely unknown . the aim of the present work was to provide a detailed account of the soft anatomy , shell microstructure , and early ontogeny of the shell and ligament in p . exempla . because this is the only living representative of the entire family and , consequently , is the only pulvinitid species known to date that can be analysed most extensively , this study provides essential data for comparative studies with other recent bivalves and establishes grounds for the interpretation of features of extinct pulvinitid taxa .
larvae at prodissoconch 1 ( p1 ) stage were isolated from the gills of an adult ( nmnz m . 150090 ) by washing the specimen in distilled water . one sample was treated with 3 % sodium hypochlorite ( 50 % dilution of commercial grade laundry bleach ) for 30 min to dissolve soft tissues , partially remove the periostracum , and allow for complete or partial separation of the valves . this and an untreated sample were transferred to the surface of stubs covered with double - sided tape in a drop of distilled water . any excess of water was carefully removed with a syringe and the remaining water was evaporated under a lamp . all sem samples were stored in a silica - gel desiccator prior to sputter coating for sem . because larval shells at prodissoconch 2 ( p2 ) stage were not available , limited observations were made on the prodissoconchs of adult and juvenile shells with the aid of light microscopy .
specimen light photography used a variety of equipment and techniques . whole - valve and detailed light micrography used a microptics® macro - imaging system based on a high - resolution nikon® single - lens reflex digital camera and a canon g2 4 . 0 mega - pixel digital camera . most histological sections were photographed using the microptics® micro - imaging system . a hamamatsu orca ii camera mounted on a leica dmra2 microscope equipped with differential interference contrast optics was used to capture images for figures 13a and 19 .
mantle edge morphology in pulvinites exempla ( ams c . 129659 , 62 . 2 mm ) . histological cross - section ( 5 µm , alcian blue / periodic acid / schiff ' s stain ) . a , overview of the mantle margin at the ventral edge of the mantle showing the proximal displacement of of - 2 ( pallial skirt ) ( scale bar = 500 µm ) . b , magnified mantle margin ( scale bar = 200 µm ) . c , close - up of of - 2 ; the arrows indicate pigmented ( presumably secretory ) cells ( scale bar = 50 µm ) . d , cross - section of the mantle edge posterior to the mantle isthmus showing the distal position of of - 2 and merging if - 1 and of - 2 folds ( scale bar = 100 µm ) . c , circumpallial artery ; if , inner mantle fold ; of , outer fold ; p , periostracum ; pgr , periostracal groove ; t , tentacle .
heart in pulvinites exempla ( ams c . 129659 , 62 . 2 mm ) . transverse histological section ( 5 µm , alcian blue / periodic acid / schiff ' s stain ) ( scale bar = 300 µm ) . au , auricles ; in , intestine ; v , ventricle .
shell measurements were taken with digital callipers from the rv . measurements on histological specimens were made using an ocular micrometer on a stereomicroscope . the maximum shell height for adult and embryonic shells was measured perpendicular to the hinge line to the most distal point of the ventral shell margin ; the maximum shell length was taken parallel to the hinge line . size is expressed as shell length unless otherwise noted . morphological and anatomical terminology follows the review by tëmkin ( 2006 ) . cited repositories include : the australian museum , sydney ( ams ) and the museum of new zealand te papa tongarewa ( nmnz ) .
hypotrema orbigny , 1853 ; foramelina hedley , 1914 [ for further information on synonymy , see palmer ( 1984 ) ] . type species by monotypy : pulvinites adansonii blainville , 1824 ; late cretaceous , france .
foramelina exempla hedley , 1914 : 71–72 , plate 11 , figure 6 , plate 12 , figures 7 , 8 ( original description ) ; – hedley , 1918 : m7 ( faunal list ) ; – thiele , 1934 : 802 ( brief description ) ; – macpherson & chapple , 1951 : 144 ( faunal list ) ; iredale & mcmichael , 1962 : 10 ( faunal list ) ; – macpherson & gabriel , 1962 : 294–295 , figure 336 ( description ) ; – cox , 1969 : n326–327 , figures c53 , 3a , b ( diagnosis of foramelina ) ; – fischer - piette , 1976 : 40 ( synonymy ) ; – tenner , 1981 : 3 .
pulvinites exempla ( hedley , 1914 ) . – palmer , 1984 : 823 , plate 72 , figure 6 ( revision ) ; – butler , 1998 : 265 , figure 6 . 11 ( description ) ; – lamprell & healy , 1998 : 110–111 , figure 267 ( brief description ) ; – marshall , 1998 : 99–102 , figures 1 – 9 ( hinge and prodissoconch figured ; ecology ) ; – malchus , 2004a : 1547 ( prodissoconch and ligament ) ; – malchus , 2004b : 96 ( prodissoconch ) ; – tëmkin , 2006 ( phylogeny ) .
pulvinites ( pulvinites ) exempla ( hedley , 1914 ) . – pacaud , 2001 : 6 , 9 , 12–14 , figure 4 ( a ) ( comparison with extinct pulvinites spp . ) .
ams c . 170923 ( one pair + byssus ) , 38°8′s , 149°55′e , australia , vic , south of gabo island , 183–457 m [ ‘type’ on hand - written label and ‘holotype’ on printed label ; ‘holotype’ in palmer ( 1984 ) and marshall ( 1998 ) ; figured in hedley ( 1914 : plate 11 , figure 6 ) ] .
ams c . 037004 ( rv ) , 38°8′s , 149°55′e , australia , vic , south of gabo island , 183–457 m [ ‘cotype’ on hand - written label and ‘paratype’ on printed label ; ‘paratype’ in palmer ( 1984 ) and marshall ( 1998 ) ; figured in hedley ( 1914 : plate 12 , figures 7 , 8 ) , macpherson & gabriel ( 1962 : 295 , figure 336 ) ; cox ( 1969 : n327 , figures c53 , 3a , b ) , palmer ( 1984 : plate 72 , figure 6 ) , butler ( 1998 : 265 , figure 6 . 11 ) , pacaud ( 2001 : 12 , figure 4a ) ] .
ams c . 122000 ( one pair ) , 33°51′s , 151°41′e , australia , new south wales , east of sydney , 400 m , trawled ; ams c . 126893 ( one pair ; rv fragment ) , 33°50 . 000′s , 151°43 . 500′e , australia , new south wales , off sydney , 457 m , trawled ; ams c . 129659 ( four pairs with tissue + one juvenile pair fragment ) , 33°42 . 000′s , 151°53 . 000′−152°0 . 000′e , australia , new south wales , north - north - east of sydney , 406 m , on a junked aeroplane , trawled ; nmnz m . 153627 ( one pair ) , 45°18 . 93′s , 166°68 . 36′e , new zealand , south island , off point east of joseph point , doubrful sound , fioedland , 340 m ; nmnz m . 150090 ( one individual tissue only + > 100
- stage larvae ) , 37°41 . 3′s , 179°00 . 7′e , off east cape , 936 m .
‘shell equivalve , equilateral , large , flat , discoidal , margin irregular in outline . valves thick , composed of brittle , imbricating lamellae which both include and are overlaid by a thin membranous epidermis , where eroded of a silvery sheen , interior a dark bronze . perforation in the right valve about 10 mm in diameter , median and subumbonal , internally with a raised margin , externally excavated as an oblique furrow ascending to the vertex . from the margin of the perforation a suture leads to the anterior extremity of the hinge plate . byssus a dense bundle of threads about half an inch long . hinge line about 43 mm long . area much eroded above and traversed by about 22 narrow ligamental grooves which slightly radiate from above . externally the hinge gapes when the valves are closed as in melina [ = isognomon solander in lightfoot , 1786 ] . adductor small , subcircular , about its own breadth below the perforation . byssal retractor rather larger than the adductor and immediately above it in the left valve . pallial margin entire , about half an inch within the ventral margin . height , 120 ; length , 130 mm’ ( hedley , 1914 : 71 ) .
‘distinguished from other species of pulvinites by its large size when adult , and by features related to size , such as the greater number of ligament pits and their greater size and spacing .
pterioidean with nongyrate and subcircular in shape prodissoconch ; anterior part of visceral mass greatly contorted to right side ; pairs of left and right anterior pedobyssal retractor muscles fused at tips ; outermost mantle fold ( of - 2 ) displaced proximally forming pallial skirt ; larvae present in gills of brooding adults .
the shell of pulvinites exempla , major features ( lectotype ; ams c . 170923 , 140 mm ) . the shells are arranged with the hinge axis orientated horizontally . a , exterior of the right valve ( rv ) . b , exterior of the left valve ( lv ) . c , interior of the lv . d , interior of the rv . e , byssus . arms , anterior ( pedobyssal ) retractor muscle scar ; cf , commarginal folds ; cl , calcitic lamellae ; cti , impressions of ctenidia ; fo , foramen ; fog , foramen groove ; hl , hinge line ; mlg , multivincular ligament ; nb , border of nacreouos layer ; pams , posterior adductor muscle scar ; pmg , prismatic margin ; pmi , impressions of pallial retractor muscles ; pms , pallial muscle scars ; prms , posterior ( pedobyssal ) retractor muscle scar ; su , suture ; u , umbo .
the diversity of shell shape in pulvinites exempla : right valve exterior . the shells are arranged with the hinge axis orientated horizontally . a , ams c . 037004 ( paralectotype ; 140 mm ) ; b , ams c . 122000 ( 90 mm ) ; c , nmnz m . 153627 ( 68 . 1 mm ) ; d , ams c . 129659 ( 45 . 2 mm ) ; e , ams c . 129659 ( 62 . 2 mm ) ; f , ams c . 129659 ( 46 . 7 mm ) . evidence of external physical damage is indicated by arrows .
juvenile specimen of pulvinites exempla ( ams c . 129659 , 3 . 91 mm ) . a , exterior of the left valve ( lv ) in lateral view , with the arrow indicating the metamorphic line . b , exterior of the right valve ( rv ; fragment ) in lateral view showing prodissoconch 2 and byssal - related structures . c , interior of the rv ( fragment ) in lateral view showing the interior of prodissoconch 2 . fog , foramen groove ; lg , resilium of ligament ; su , suture .
prodissoconch 2 ( p2 ) of the left valve of pulvinites exempla ( juvenile , ams c . 129659 , 3 . 91 mm ) . the orientation of p2 is given with regard to its placement in the dissoconch . a , dorsal view of the exterior showing the concentric sculpture of p2 and its opisthogyrate placement . the white arrows indicate the p1–p2 transition ; the black arrow indicates the metamorphic line . the hinge axis is orientated horizontally with the anterior at the left . b , ventral view showing the interior surface of p2 and the initial crenulation of the hinge plate underlying the first resilium . the arrow indicates the border of confluent scars of the posterior adductor–posterior pedobyssal retractor muscles . c , lateral view of the exterior surface showing the extent of inflation of p2 . the white arrow indicates the p1–p2 transition ; the black arrow indicates the metamorphic line . lg , resilium of ligament .
in addition to the muscle scars deeply impressed in the shell interior , some shells bear clear impressions of the gills and pallial retractor muscles ( obvious in large thick - shelled specimens ) ( fig . 1c , d ) . some specimens bear a sign of external physical damage affecting one or both valves , but subsequent shell deposition shows a successful recovery ( fig . 2b , c , f ) .
( scanning electron microscopy ; nmnz m . 150090 ) . a , overview of the interior of joined valves in lateral view showing the
- shaped outline and straight hinge ( scale bar = 100 µm ) . b , exterior surface of a valve in lateral view ; the close - up of the squared area ( inset ) shows a nodulose surface ; the arrow indicates the commarginal fold demarcating the marginal rim ( scale bar = 100 µm ) . c , exterior surface of a valve in dorsal view showing faint commarginal and radial striae ; the arrow indicates the periostracal ligament ( scale bar = 10 µm ) . d , lateral view of the inner surface showing the granular interior surface and smooth valve margin ( scale bar = 2 µm ) .
morphology of the provinculum in prodissoconch 1 of pulvinites exempla ( scanning electron microscopy ; nmnz m . 150090 ) . a , interior view of the hinge plate of an articulated shell ; the inset lettering refers to magnified views of outlined areas in corresponding figures ( scale bar = 50 µm ) . b , transverse denticles ( scale bar = 2 µm ) . c , d , terminal teeth and corresponding sockets ( scale bar = 10 µm ) . the arrow indicates the periostracal ligament .
a large sample of the ontogenetically earliest shell - p1–was obtained from the gills of a large individual ( nmnz m . 150090 ) . p1 ranges from 289 to 299 µm in length ( mean 292 ± 3 . 54 µm ,
) . the valves are approximately equiconvex and moderately inflated , producing an oblong flattened globe . the ‘punctate–stellate pattern’ of the p1 shell surface previously reported in ostreids (
) was not observed . instead , the exterior surface is smooth but at higher magnification appears to have multiple low nodules , which are slightly elongated with the long side orientated commarginally (
, inset ) . these structures are distributed irregularly across most of the valve ' s surface , except for the area along the hinge line and the medial area extending ventrally from the hinge to approximately one - third of the valve height . very faint , irregular , and tightly packed commarginal striae and more broadly spaced irregular radial striae are present on the smooth part of the exterior surface (
) . the inner surface is granular . the margins of p1 are not interlocking , but are thickened and smooth ( relative to the inner surface ) , ensuring a precise tight closure (
well - defined interlocking provincular teeth are present on the anterior and posterior ends of the hinge line in the area where it is continuous with the anterior and posterior margins ( fig . 5 ) . thus , the teeth are terminal and arranged on a slight curve in a widening flat area marking the origin of the marginal rim . the teeth are symmetrical between the valves and are typically three at the anterior and posterior end of each valve , whereas two specimens were observed with two teeth and one with four ( n = 10 pairs ) . the teeth of a single row are approximately the same height , but vary in width : some teeth are twice as wide as neighbouring ones . the narrow hinge plate between the anterior and posterior teeth is covered by minute transverse and poorly defined denticles that potentially provide additional reinforcement against a shearing force ( fig . 5b ) .
p1 is distinguished from the veliger shell - p2–by the first commarginal growth line and the presence of distinct commarginal sculpture in the latter ( fig . 6a , c ) . p2 ranges from 550 to 616 µm in length ( mean 583 , n = 2 ) . a prominent umbo is lacking , resulting in the rounded outline of p2 . the shell is , however , inequilateral due to the posterior coiling of p2 and ultimately becomes opisthogyrate , expanding along the anteroventral margin . both valves are considerably convex . the exterior surface of p2 has conspicuous , close - set , regularly spaced commarginal growth lines . the interior surface of the valves is smooth at low magnification . the present study confirmed previous observations of the external morphology of a fully formed larval shell by marshall ( 1998 ) .
the fate of the provincular teeth could not be traced in the p2 stage , due to the unavailability of free shells at that stage . however , by analogy with ostreid taxa in which the ontogeny of provincular teeth has been described in considerable detail ( carriker & palmer , 1979 ; waller , 1981 ) , the general arrangement of the teeth probably persists in the p2 stage , but subsequently the teeth become obliterated by additional deposition of shell material later in ontogeny . for the same reason , the state of marginal characters in p2 , such as the presence of marginal denticles and tongue - in - groove interlocking margins , could not be verified . however , a single available early juvenile ( ams c . 129659 , 3 . 91 mm ; figs 3 , 6 ) shell allowed limited observations on the exterior and interior surfaces of p2 . no shell septum was observed on the interior surface . the exterior of the lv of p2 as seen in the juvenile and adult shells does not show evidence of the fasciole and the associated posterodorsal notch and outlet .
p2 is sharply demarcated from the early dissoconch by a metamorphic line ( figs 3a , 6a , c ) . postmetamorphic growth greatly affects the relative position of the left and right p2 . in an early juvenile ( one resilifer stage ) , the left p2 is placed nearly horizontally dome - like in a more inflated lv , whereas the right p2 is placed vertically in the rv at nearly a right angle to the left p2 . in the adult , the larval shells are widely separated by the space between the hinge plates and the left p2 rotates even further dorsally around the larval hinge line .
the large size of p1 ( > 100 µm ) , the high p1 / p2 ratio ( 0 . 5 ) , and the absence of a prominent larval umbo indicate a lecithotrophic–planktotrophic reproductive mode and are often correlated with brooding behaviour [ reviewed by malchus ( 2004a ) ] .
mineralogically , p1 and p2 are presumably composed of homogeneous aragonite , as suggested by previous observations in other related pteriomorphian taxa : pinctada fucata martensii ( dunker , 1872 ) by kobayashi ( 1980 ) and in oysters by stenzel ( 1964 ) and carriker & palmer ( 1979 ) . the aragonitic composition of larval shell appears to be uniform across the bivalvia ( stenzel , 1964 ; taylor , kennedy & hall , 1969 ) .
outer shell layer in pulvinites exempla ( scanning electron microscopy ; ams c . 129659 , 45 . 2 mm ) . a , cross - section of shell showing the inner nacreous ( top ) and outer prismatic ( bottom ) layers ( scale bar = 150 µm ) . the arrow indicates the layer of epoxy that filled the space between separated shell layers . b , exterior surface of the outer shell layer covered with periostracum ( scale bar = 200 µm ) . c , close - up lateral view of prisms showing interprismatic conchiolin walls ( black arrows ) and accretion lines ( white arrows ) ( scale bar = 25 µm ) . d , close - up of the exterior surface of the outer shell layer showing the worn periostracum and the exposed dorsal surface of prisms ( scale bar = 50 µm ) . the arrows indicate interprismatic junctions . e , exterior surface of prisms in lateral view overlain by a periostracal layer ( p ) ( scale bar = 10 µm ) . f , dorsal view of the outer shell surface showing the cross - sectional shape of prisms and the partially abraded periostracum ( p ) ( scale bar = 25 µm ) . g , lamella of the outer surface ( arrow ) ( scale bar = 50 µm ) . h , pockets of the prismatic layer ( arrows ) ( scale bar = 100 µm ) . ( a ) , ( c ) , ( g ) , and ( h ) were prepared by polishing and etching sections of epoxy - embedded left valve ; ( b ) , ( d ) , and ( e ) were prepared by vertical mechanical fracture of a right valve ; ( f ) was prepared by vertical mechanical fracture and subsequent bleaching treatment of a right valve .
nacre in pulvinites exempla ( scanning electron microscopy ; ams c . 129659 , 45 . 2 mm ) . a , cross - section of shell showing the inner nacreous layer containing irregular pockets ( arrows ) ( scale bar = 10 µm ) . b , magnified lateral view of nacreous tablets showing the intercrystalline organic matrix ( white arrows ) and the tuberculate surface ( scale bar = 3 µm ) . c , rhomboid tablets of the outer surface of the nacreous layer ( scale bar = 20 µm ) . inset : a magnified single tablet showing the tuberculate surface ( scale bar = 1 µm ) . d , magnified surface of a nacre tablet showing sinistral spiral growth ( scale bar = 2 . 5 µm ) . ( a ) and ( b ) were prepared by polishing and etching sections of epoxy - embedded left valve ; ( c ) and ( d ) were prepared by vertical mechanical fracture of a right valve .
myostracum in pulvinites exempla ( scanning electron microscopy ; ams c . 129659 , 45 . 2 mm ) . a , cross - section of shell showing the myostracum separating the inner and outer nacreous layers ( scale bar = 100 µm ) . b , magnified cross - section of the interface of the myostracum and the outer nacreous layer showing the orientation of the irregular prismatic myostracum ( scale bar = 5 µm ) . m , myostracum ; n i , inner nacreous layer ; n o , outer nacreous layer ; p , prismatic layer . the figures were prepared by polishing and etching sections of epoxy - embedded left valve .
the inner layers are sheet nacreous with tablets reaching 1 . 2 µm in thickness ( fig . 8 ) . the deposition of nacre is not uniform and irregular layers of six to ten tablets are formed by narrow empty pockets within the nacreous structure . individual tablets are very regularly rhomboid in shape and greatly vary in size depending on the growth stage . the predominant mode of growth of the nacreous layer is by nucleation of small crystals with their subsequent coalescence within the same layer , separated from an older layer by a thick interlamellar organic matrix . individual tablets at the outermost layer of the inner shell surface are distantly and irregularly spaced , but later become aligned along parallel faces . as they increase in size , the crystals form a dendritic layer connecting with neighbouring crystals by intercrystalline organic matrix . at the terminal stage of growth , a continuous sheet of nacre is formed with intercrystalline junctions concealed by an overlaying interlamellar organic matrix . at that stage , the regular rhomboid outline is modified to accommodate a tight contact with neighbouring tablets . in addition , some tablets display dextral or sinistral spiral surface growth with a polygonal outline ( fig . 8d ) .
a continuous layer of the adductor myostracum separates the inner and middle nacreous layers and reaches 120 µm thickness at the point of its emergence on the inner shell surface ( fig . 9 ) . microstructurally , it consists of highly irregular ( in size and outline ) prisms that are orientated strictly perpendicular to the underlying sheets of nacre . mineralogically , the myostracum is assumed to be aragonitic , as shown to be invariable in bivalves ( taylor et al . , 1969 ) .
ligament morphology in pulvinites exempla . a , lateral view of the left valve hinge plate showing the orientation of the ligament relative to the umbo ( arrow ) and the relative position of fibrous ( f ) and lamellar ( l ) sublayers . superscripts indicate the ontogenetic order of the deposition of resilia ( ams c . 129659 ; 62 . 2 mm ) . b , lateroventral view of the left valve hinge plate showing the breached ventral edge and the rounded cross - section of resilifers . note the difference in colour between the ligostracal resilifer walls and the underlying ostracum ( nmnz m . 153627 ; 68 . 1 mm ) .
ligament microstructure in pulvinites exempla ( scanning electron microscopy ; ams c . 129659 , 45 . 2 mm ) . a , transverse section of the hinge plate made through a resilifer showing the arrangement of shell layers ( scale bar = 1 mm ) . enlargement of the squared area is presented in ( b ) . b , widest area of the ligostracum making the roof of the resilifer ( scale bar = 50 µm ) . c , magnified interface of the ligostracal and nacreous layer showing the irregular pattern of the ligostracum ( scale bar = 15 µm ) . flg , fibrous ligamental sublayer ; l , ligostracum ; m , myostracum ; n i , inner nacreous layer ; n o , outer nacreous layer ; p , prismatic layer . figures were prepared by polishing and etching sections of epoxy - embedded left valve .
the hinge plate is a triangular thickened area located in the middle of the hinge line with the prodissoconch situated at the dorsalmost extremity ( fig . 10a ) . the hinge plate progressively expands in width from occupying approximately one - third of the hinge line ' s length in juveniles to nearly its entire length in adults . the base ( ventral edge ) of the hinge plate is slightly produced towards the opposite valve , resulting in inclination of its surface , which is stronger in the rv . such a position of the hinge results in a v - shaped commissure ( when viewed end on ) of the dorsal margins allowing for opening of the valves . the hinge plate is excavated below its ventral margins in both valves and slightly crenulated on the back side .
the pattern of resilifer origination suggests an alternating order of appearance of anterior and posterior resilifers ( relative to the oldest one ) , as previously suggested by malchus ( 2004a ) . the rate of insertion of anterior resilifers is , however , higher than that of the posterior resilifers . the highest ( and presumably oldest ) resilifer is initially directed slightly anteriorly and originates ventral to the prodissoconch , suggesting that it might represent the continuation of the larval ligament . this interpretation is corroborated by observation of the ligament in the only available early juvenile specimen . at this stage ( fig . 3c ) , only one submarginal resilium is present and is directed anteroventrally , originating from approximately the centre of the provincular hinge line . the resilifer is not formed yet , but the ventrally expanding margin of the resilium is supported by a thickened and weakly crenulated larval hinge plate . the lamellar component is visible , stretching along the provincular axis anteriorly and posteriorly to the resilium . thus , examples of the ligament growth formula ( malchus , 2004a ) for p . exempla are 7p - v , 4p - v / 1c - v , 2a - v , 3a - v , 5a - v , 6a - v , 8a - v ( ams c . 129659 ; 62 . 2 mm ) ; 6p - v , 4p - v / 1c - v , 2a - v , 3a - v , 5a - v , 7a - v ( ams c . 129659 ; 46 . 7 mm ) . it must be added that the initial growth direction of the first resilifer is towards the anterior .
the surface of the hinge plate is coated with a thin , irregularly structured , and sponge - like layer of ligostracum separating the ligament from adjacent shell layers ( fig . 11 ) . the walls of the resilifers consist entirely of ligostracum . ligostracum is approximately 15 µm thick along most of its length , but slightly widens at the ventralmost extremity of the resilium to 35 µm in thickness . it is most extensive at the area dorsal to the resilium , where it forms a roof of the resilifer ( reaching up to 120 µm thickness ) ( fig . 11b ) .
the overall body plan of p . exempla ( fig . 12 ) displays dramatic contortion of the anterior part of the body , probably a result of adaptation to a pleurothetic mode of life with the byssus projecting through the foramen in the rv . the visceral mass is very small relative to the size of the shell , and is restricted to the small area underneath the hinge . it is entirely concealed by the byssus in lateral view once the rv is removed ( fig . 12b ) . the visceral mass is produced posteroventrally towards the anteroventral surface of the pam and is connected to it by very loose connective tissue . in addition to the extreme contortion , the anteroposterior axis [ as defined in jackson ( 1890 ) ] of the visceral mass is at an acute angle to the hinge line . the orientation of structures in the following discussion of the soft anatomy is relative to the anteroposterior axis .
overview of the soft anatomy of pulvinites exempla ( ams c . 129659 , 62 . 2 mm ) . a , right lateral view of the soft anatomy with the valves , byssus , right mantle lobe , and right ctenidium removed . b , right lateral view of the intact soft anatomy with the right valve removed . c , left lateral view of the soft anatomy with the byssus , valves , right mantle lobe , and right ctenidium removed . d , diagrammatic view of the soft anatomy showing the alimentary tract and musculature . e , diagrammatic posterior view of transverse section through midline showing relative arrangement of principle muscles . arm , anterior pedobyssal retractor muscle ; by , byssus ( cut ) ; byf , byssal fissure ; ct , ctenidium ; f , foot ; h , heart ; in , intestine ; k , kidney ; lg , ligament ; li , lips ; lp , labial palp ; lv , left valve ; mi , mantle isthmus ; ml , mantle lobe ; mo , mouth ; oe , oesophagus ; pam , posterior adductor muscle ; pm , pallial muscles ; prm , posterior pedobyssal retractor muscle ; psk , pallial skirt ; rv , right valve ; sm , suspensory membrane ; ss , supramyal septum ; st , stomach ; vg , visceral ganglia ; vm , visceral mass .
details of mantle edge morphology in pulvinites exempla ( scanning electron microscopy ; ams c . 129659 , 62 . 2 mm ) . a , cross - section of the mantle ; the arrow shows of - 2 ( pallial skirt ) ( scale bar = 200 µm ) . b , lateral view of the interior mantle margin showing the outer surfaces of if - 1 , if - 2 , and of - 1 ; the arrow indicates the sinuous groove separating the inner fold ( scale bar = 200 µm ) . c , magnified large tentacle of the inner surface of if - 1 ( scale bar = 25 µm ) . d , small laterally compressed tentacles of the outer surface of if - 2 ( scale bar = 50 µm ) . e , ciliated inner surface of if - 1 ( scale bar = 10 µm ) . f , smooth inner surface of of - 1 ( scale bar = 10 µm ) ; if , inner mantle fold ; of , outer fold ; p , periostracum ; t , tentacle .
the mantle lobes occupy the entire area between the valves , extending from the mantle isthmus around the entire circumference of the shell margin . relative to the size of the visceral mass , the extent of the pallial cavity is very large ( fig . 12a ) . the margins of the mantle lobes are fused dorsally immediately under the hinge , forming a mantle isthmus with a regularly crenulated outline . the crenulations of the mantle isthmus correspond to the most proximal ( newly secreted ) , ventrally protruding extremities of the resilia . the area of mantle fusion extends slightly anterior and posterior to the ligament and apart from this restricted area along the hinge line , the margins of the mantle lobes are free . the inner surfaces of the left and right mantle lobes are connected by a supramyal septum , a thin cross membrane , stretching from the posterodorsal surface of the adductor muscle to the mantle isthmus in the proximity of the posterior fusion of mantle margins ( fig . 12a , c ) . a short stretch of tissue connects the septum with the posterodorsal surface of the visceral mass and the left mantle lobe , producing a blind pocket on the left side of this chamber slightly ventral to the posterior section of the hinge line . a fissure , present in the right mantle lobe , extends posteroventrally at an acute angle from the area slightly anterior to the mantle isthmus to the area of the emergence of the byssus ( fig . 12d ) . the byssal fissure is narrow along its length with the exception of the widening accommodating the passage of the byssus . the fissure is mirrored in the rv by the ontogenetic suture .
the inner surfaces of the mantle lobes also form connections to the muscles and the viscera . the only symmetrical area of mantle attachment on both sides includes the pam that perforates the mantle lobes . the inner surface of the right mantle lobe is fused along a narrow area produced along the proximal edge of the outer labial palp to the anterodorsal part of the adductor muscle , passing through the attachment area of the distal extremities of the right pair of arms . the left mantle lobe is partially fused to the outer surface of the upper lip , the outer surface of the left pair of arms , and the prm that perforates the lobe .
the mantle edge is divided into four folds ( fig . 13 ) . the periostracal groove separates the two inner folds from the two outer folds . the middle fold ( if - 1 ) is very short ; its outer surface forms the inner wall of the periostracal groove and the inner surface bears irregularly spaced , short , stout , round in cross - section , finger - like ciliated tentacles ( figs 13b , 14b , c ) . some of these tentacles have first - order branches of approximately the same thickness . the inner mantle fold ( if - 2 ) is separated from if - 1 by a wavy groove . the inner and outer surfaces of if - 2 have commarginal grooves and folds . the outer surface of if - 2 is equipped with short irregularly spaced unbranched tentacles , somewhat smaller than those of if - 1 and laterally compressed , extending along the commarginal grooves ( fig . 14b , d ) . bundles of pallial muscles extend to the edge of if - 2 along its inner surface . the surfaces of if - 1 and if - 2 are ciliated ( fig . 14c–e ) , as are the inner and outer surfaces of distal mantle margins . periostracum is secreted from a cleft between of - 1 and if - 1 ( figs 13b , d , 14b ) .
the outer fold proximal to the periostracal groove ( of - 1 ) is most extensive . it is covered by an extremely thin layer of smooth epithelium ( underlain by very loose tissue ) on both sides and is devoid of secreting cells ( with the exception of pigmented large cells forming the outer wall of the periostracal groove ) . no tentacles , nerves , or muscular bundles were detected in its interior . of - 1 does not maintain a regular outline . the outermost fold ( of - 2 ) is removed from the mantle margin proximally , forming a conspicuous commarginal pallial skirt at approximately 2 . 5 mm from the mantle edge ( figs 12a , 13a , c , 14a ) . as it approaches the mantle isthmus , of - 2 is gradually displaced distally to a position immediately adjacent to of - 1 ( fig . 13d ) . in this dorsal region , all mantle folds reach nearly the same height . pallial muscles stretching close to the outer surface of the mantle lobes are produced into of - 2 and terminate at this point ; only loose muscle fibres are produced distally of this point . of - 2 contains pigmented granules , suggestive of its role in secretion . pigmented cells are also present at a very low concentration on if - 2 ( fig . 13c ) and the adjacent portion of the inner mantle surface . the remaining mantle margin and the surfaces of the mantle lobes lack pigmentation and are effectively translucent .
a cilial trail , along which a cilial connection is made between the mantle and outer demibranchs , stretches medially on the inner surface of the mantle lobes from an area close to the ventral edges of the labial palps to the mid - posterior edge of the mantle . if - 2 is thickened at the point of contact of the distalmost extremity of the cilial path and the inner surface of the mantle edge .
posterior adductor muscle in pulvinites exempla ( ams c . 129659 , 62 . 2 mm ) . histological sagittal section ( 5 µm , alcian blue / periodic acid / schiff ' s stain ) . the section was made close to the attachment surface of the adductor muscle showing slightly denser packed muscle fibres in the anterior section ( right side ) than in the posterior section ( left side ) , separated by the dashed line ( scale bar = 150 µm ) . inset : visceral transverse connective ( vc ) suspended in loose connective tissue at the anteroventral surface of the posterior adductor muscle ( pam ) ( scale bar = 250 µm ) .
the most conspicuous muscle is a hypertrophied left prm that extends from the left side of the visceral mass ( in a direction opposite to that of the byssus ) , producing a large circular muscle scar a short distance below the hinge line in the lv ( fig . 12c–e ) . the muscle is composed of radially striated muscle bundles . a corresponding right prm is absent . a reduced , circular in cross - section , pam forms a pair of round symmetrical scars in each valve , located ventrally either directly under or slightly posterior to the posterior retractor muscle scar in the lv and the foramen in the rv ( fig . 12 ) . the anterior adductor is absent ( monomyarian condition ) . despite the fact that the surface of the adductor muscle does appear to consist of two ( slightly different in texture ) subequal anterior ( ‘quick’ ) and posterior ( ‘catch’ ) portions separated by an irregular wavy line , the fibres of the two parts are of the same thickness and are indistinguishable at the microscopic level when viewed in cross - section through the middle of the muscle . closer to its left and right surface , the fibres of the posterior side appear to be more loosely packed than those of the anterior side , which might explain the pattern seen on the exterior . the adductor muscle is homogeneous and comprised exclusively of loosely arranged opaque transverse ‘catch’ ( ‘slow’ ) smooth fibres ( by comparison with ‘catch’ muscles of other species ; kennedy , newell & eble , 1996 ; mikkelsen et al . , 2004 ) ( fig . 15 ) .


doleromyrma is a genus of ants in the subfamily dolichoderinae .

the genus is native to australia , where the ants nest in soil , or under stones or logs .

doleromyrma

the following species and subspecies belong to the genus doleromyrma . synonyms are listed under their senior names . for valid names only see doleromyrma species and for distribution information see doleromyrma species by country .
richness of doleromyrma species ( countries with darker colours are more species - rich ) . for a list of species and subspecies see the checklist of doleromyrma species or for valid names only see doleromyrma species .
doleromyrma forel , 1907a : 28 [ as subgenus of tapinoma ] . type - species : tapinoma ( doleromyrma ) darwinianum , by monotypy .
the above specimen data are provided by antweb . please see doleromyrma darwiniana for further details
doleromyrma in dolichoderinae , leptomyrmecini : ward , brady , et al . 2010 : 361 .
doleromyrma as subgenus of tapinoma : forel , 1907d pdf : 28 ; wheeler , 1910a pdf : 142 .
doleromyrma revived from synonymy and raised to genus : shattuck , 1992a : 14 ; shattuck , 1992c : 60 .
bolton , b . 1994 . identification guide to the ant genera of the world . cambridge , mass . : harvard university press , 222 pp . ( page 26 , doleromyrma as genus ; doleromyrma in dolichoderinae , dolichoderini )
combination in iridomyrmex : emery , 1913a pdf : 24 ; in doleromyrma : shattuck , 1992a pdf : 14 .
doleromyrma junior synonym of iridomyrmex : emery , 1913a : 22 ; wheeler , w . m . 1922a : 689 .
doleromyrma as genus : shattuck , 1992a pdf : 14 ; bolton , 1994 : 26 ; bolton , 2003 pdf : 84 .
doleromyrma in dolichoderinae , dolichoderini : shattuck , 1992c pdf : 60 ; bolton , 1994 : 26 ; bolton , 2003 pdf : 84 .
doleromyrma in dolichoderinae , tapinomini : emery , 1913a pdf : 22 ; donisthorpe , 1943g : 640 ; hölldobler & wilson , 1990 : 17 .
doleromyrma as junior synonym of iridomyrmex : emery , 1913a pdf : 22 ; wheeler , 1922 : 689 ; all subsequent authors to the following .
bolton , b . 2003 . synopsis and classification of formicidae . mem . am . entomol . inst . 71 : 370pp ( page 84 , doleromyrma as genus )
tapinoma ( doleromyrma ) darwinianum forel , 1907 : syntype , 2 workers , mt . victoria , blue mountains , new south wales , australia , museum of comparative zoology .
forel , a . 1912c . einige interessante ameisen des deutschen entomologischen museums zu berlin - dahlem . entomol . mitt . 1 : 81 - 83 ( page 81 , doleromyrma subgenus of iridomyrmex )
shattuck , s . o . 1992c . generic revision of the ant subfamily dolichoderinae ( hymenoptera : formicidae ) . sociobiology 21 : 1 - 181 ( page 60 , doleromyrma in dolichoderinae , dolichoderini )
tapinoma ( doleromyrma ) darwinianum forel , 1907 : syntype , 3 workers , 1 male , mt . victoria , blue mountains , new south wales , australia , musee d ' histoire naturelle genève .
wheeler , w . m . 1910b . ants : their structure , development and behavior . new york : columbia university press , xxv + 663 pp . ( page 142 , doleromyrma as subgenus of tapinoma )
forel , a . 1907d . formicides du musée national hongrois . ann . hist . - nat . mus . natl . hung . 5 : 1 - 42 ( page 28 , doleromyrma as subgenus of tapinoma )
hölldobler , b . ; wilson , e . o . 1990 . the ants . cambridge , mass . : harvard university press , xii + 732 pp . ( page 17 , doleromyrma in dolichoderinae , tapinomini )
tapinoma ( doleromyrma ) darwinianum forel , 1907 : syntype , 1 worker , mt . victoria , blue mountains , new south wales , australia , < collector unknown > , anic32 - 001666 , australian national insect collection .
shattuck , s . o . 1992a . review of the dolichoderine ant genus iridomyrmex mayr with descriptions of three new genera ( hymenoptera : formicidae ) . j . aust . entomol . soc . 31 : 13 - 18 ( page 14 , combination in doleromyrma )
donisthorpe , h . 1943g . a list of the type - species of the genera and subgenera of the formicidae . [ part ] . ann . mag . nat . hist . 11 ( 10 ) : 617 - 688 ( page 640 , doleromyrma in dolichoderinae , tapinomini )
shattuck , s . o . 1992a . review of the dolichoderine ant genus iridomyrmex mayr with descriptions of three new genera ( hymenoptera : formicidae ) . j . aust . entomol . soc . 31 : 13 - 18 ( page 14 , doleromyrma revived from synonymy with status as genus )
workers in the genus doleromyrma have 12 - segmented antennae . the mandibles have 4 - 5 teeth and 4 - 5 denticles . the front margin of the clypeus has downwardly curved hairs . the upper face of the propodeum is shorter than the rear face . the petiolar node is inclined forward .
doleromyrma contains a single described species and two
subspecies
and has received little attention in the literature . it was originally established as a subgenus of tapinoma ( forel 1907 ) , was then synonymised with iridomyrmex ( emery 1912 ) , but is now treated as a distinct genus ( shattuck 1992 ) .
wheeler , w . m . 1922i . ants of the american museum congo expedition . a contribution to the myrmecology of africa . vii . keys to the genera and subgenera of ants . bull . am . mus . nat . hist . 45 : 631 - 710 ( page 689 , doleromyrma junior synonym of iridomyrmex )
apart from some spreading northward in the auckland area , this species still tends to remain associated very closely with towns or cities with ports , strongly suggesting separate port invasions in some cases . thus it has been recorded in whangarei , mt maunganui , gisborne , napier , blenheim , nelson and lyttelton .
length about 2 mm ; colour of head dark brown , rest of the body and legs light brown . refer also to the generic features above . similar in appearance to argentine ants ( l . humile ) but workers are easily separated in the field as they give off a strong odour when crushed ( cf . little or no odour for argentine ants ) .
taylor ( 1959 ) noted numerous portions of arthropod exoskeleton in the nest referred to above . this species is clearly a generalist because he also observed a small group of workers tending root - feeding homoptera ; and in the laboratory , sugar and honey were
eagerly accepted .
appear to spread predominantly by budding and can build up large densities . commonly found nesting in situations such as potted plants , probably facilitating their spread around new zealand .
not capable of stinging but will occasionally enter houses in large numbers foraging for sweet foods . attains large densities in urban gardens becoming a nuisance and may displace other invertebrates . tends aphids and mealy bugs and may also spread disease ( keall 1979 ) . has not been collected from native habitats .
the front margin of the clypeus above the mandibles with downwardly curved hairs which are about one - half the length of the closed mandibles . the mandibles with 4 or 5 large teeth and 4 or 5 small denticles , and with a moderately distinct angle between the surface containing the teeth and the surface near the clypeus ( basal angle present ) . the upper face of the propodeum is shorter than the rear face . the petiole is strongly inclined forward but is angular above and the forward and rear faces are present and distinct . the first segment of the gaster projects forward and partially conceals the petiole when viewed from above .
species richness by country based on regional taxon lists ( countries with darker colours are more species - rich ) . view data
the following information is derived from barry bolton ' s new general catalogue , a catalogue of the world ' s ants .
unless otherwise noted the text for the remainder of this section is reported from the publication that includes the original description .
shattuck , s . o . 1999 . australian ants . their biology and identification . collingwood , victoria : csiro publishing , xi + 226 pp . ( page 67 , australia synopsis )
this page was last modified on 26 august 2016 , at 23 : 20 .
forel , 1907a : 28 ( w . m . ) australia . imai , crozier & taylor , 1977 : 345 ( k . ) . combination in
forel , a . 1907d . formicides du musée national hongrois . ann . hist . - nat . mus . natl . hung . 5 : 1 - 42 ( page 28 , worker , male described )
imai , h . t . ; crozier , r . h . ; taylor , r . w . 1977 . karyotype evolution in australian ants . chromosoma ( berl . ) 59 : 341 - 393 ( page 345 , karyotype described )
this page was last modified on 17 may 2018 , at 23 : 23 .
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forel , a . , 1912 , h . sauter ' s formosa - ausbeute : formicidae ( hym . ) . , entomologische mitteilungen 1 , pp . 45 - 81
als ich diese untergattung aufstellte ( termeszetrajzi fuezetek 1907 , p . 28 - 29 ) , wusste ich nicht , ob ich sie zu
, da der kaumagen ( pumpmagen ) nicht untersucht werden konnte . seither erhielt ich eine neue varietaet (
) der gleichen art , ver - saeumte aber bisher die untersuchung des kaumagens vorzunehmen . . nun teilte mir neulich mein freund prof . emery mit , er neige eher dazu , der schuppe wegen , diese untergattung zu
hat . ich verweise auf fig . 5 und 6 der pl . xxiii , vol . xv des bulletin de la soc . vau - doise des sciences naturelles 1878 : forel etudes myrmecologiquos en 1878 . - diese figuren passen ziemlich genau auch fuer dolero - myrma , so dass ich nun die untergattung zu
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from the crazylegs creations range of hand crafted hebel formicariums comes a nest with a very unique appearance .
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white soil times found in banksia woodland , white sand times found in heathland , 0 times found in lateritic soil with granite outcrops . dry sclerophyll woodland with borya nitida on and around outcrops , 1 times found in mixed native / exotic veg . , vegetated coastal dune .
imai , crozier & taylor , 1977 pdf : 345 ( k . ) .
1 times found in eucalypt woodland , 1 times found in dry sclerophyl , 0 times found in dry sclerophyll .
1 times under stone , 0 times nest under rock , 1 times nest in soil under stone .
shattuck , s . o . 1992 ,
generic revision of the ant subfamily dolichoderinae ( hymenoptera : formicidae )
, sociobiology , vol . 21 , pp . 1 - 181
scott , r . r . ; emberson , r . m . 1999 : handbook of the new zealand insect names : common and scientific names for insects and alllied organisms . bulletin of the entomological society of new zealand 12 : 100pp
urn : lsid : biodiversity . org . au : afd . taxon : 0fab7be5 - a880 - 4464 - 915b - d29de2e7f75c
urn : lsid : biodiversity . org . au : afd . taxon : 29bcfe4f - 32a2 - 4cec - 8f73 - dbf8bfe2cea5
urn : lsid : biodiversity . org . au : afd . taxon : 3f79a5b4 - 8898 - 4cee - b091 - 55a2fc2d5acd
urn : lsid : biodiversity . org . au : afd . taxon : 47a09752 - 8848 - 4cdc - a8ee - db872c9d3d2e
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