title: Behavioural Endocrinology of Breeding Adelie Penguins (Pygoscelis adeliae)

authors: Ninnes, Calum Edward

subjects: Adelie penguin

abstract: Measuring hormonal changes is vital for understanding how the social and physical environment influences behaviour, reproduction and survival. Various methods of hormone measurement exist, potentially explaining variation in results across studies; methods should be cross validated to ensure they correlate. I directly compare faecal and plasma hormone measurements (Chapter 2), and use the most suitable endocrine measure to test the Darling hypothesis (Chapter 3) - that breeding is hastened and synchronized in larger colonies due to increased social stimulation (mediated by the endocrine system).
Blood and faecal samples were simultaneously collected from individual Adelie penguins (Pygoscelis adeliae) for comparison, and assayed for testosterone and corticosterone (or their metabolites). Sex differences and variability within each measure, and correlation of values across measures were compared. For both hormones, plasma samples showed greater variation than faecal samples. Males had higher corticosterone levels than females, but the difference was only significant in faecal samples. Plasma testosterone, but not faecal testosterone, was significantly higher in males than females. Correlation between sample types was poor overall, and weaker in females than in males; perhaps because measures from plasma represent hormones that are both free and bound to globulins, whereas measures from faeces represent only the free portion. Faecal samples also represent a cumulative measure of hormones over time, as opposed to a plasma 'snapshot' concentration. Faecal sampling appears more suitable for assessing baseline hormone levels.
In the second study I examined, over two seasons, whether the timing of breeding varied with colony size; larger colonies present occupants with higher levels of social stimulation and are predicted to show earlier, more synchronous breeding. Baseline faecal hormone levels throughout the breeding season, and survival, were measured to investigate possible proximate and ultimate mechanisms for the results. The influence of environmental variability was examined, by relating the timing of breeding, survival, and endocrine changes to sea ice conditions. Colony size did not influence the timing or synchrony of breeding, survival, or hormone levels within years; perhaps because colonies in an Adelie rookery are not independent from the 'social environment' of adjacent colonies. Across years, synchrony in the smaller rookery was higher than in the larger rookery. The scale of these comparisons may exceed the applicability of the Darling hypothesis. Therefore, no support was found for the Darling hypothesis, at the colony or rookery level, in this species. Higher corticosterone metabolite and lower sex hormone levels in the first season correlated to later breeding and lower survival compared to the second season. This is likely due to the persistence of extensive sea ice conditions late into the first season.
Researchers should take care in selecting the most appropriate method of hormone measurement for their question. Future studies testing the Darling hypothesis must carefully select their definition of a colony (i.e. a truly isolated social unit) and the scale at which the hypothesis is tested. Combining endocrine measurements with behavioural, survival, and environmental information allows for a more comprehensive interpretation of animal ecology.

text: Behavioural Endocrinology of Breeding Adelie Penguins (Pygoscelis adeliae) Measuring hormonal changes is vital for understanding how the social and physical environment influences behaviour, reproduction and survival. Various methods of hormone measurement exist, potentially explaining variation in results across studies; methods should be cross validated to ensure they correlate. I directly compare faecal and plasma hormone measurements (Chapter 2), and use the most suitable endocrine measure to test the Darling hypothesis (Chapter 3) - that breeding is hastened and synchronized in larger colonies due to increased social stimulation (mediated by the endocrine system).
Blood and faecal samples were simultaneously collected from individual Adelie penguins (Pygoscelis adeliae) for comparison, and assayed for testosterone and corticosterone (or their metabolites). Sex differences and variability within each measure, and correlation of values across measures were compared. For both hormones, plasma samples showed greater variation than faecal samples. Males had higher corticosterone levels than females, but the difference was only significant in faecal samples. Plasma testosterone, but not faecal testosterone, was significantly higher in males than females. Correlation between sample types was poor overall, and weaker in females than in males; perhaps because measures from plasma represent hormones that are both free and bound to globulins, whereas measures from faeces represent only the free portion. Faecal samples also represent a cumulative measure of hormones over time, as opposed to a plasma 'snapshot' concentration. Faecal sampling appears more suitable for assessing baseline hormone levels.
In the second study I examined, over two seasons, whether the timing of breeding varied with colony size; larger colonies present occupants with higher levels of social stimulation and are predicted to show earlier, more synchronous breeding. Baseline faecal hormone levels throughout the breeding season, and survival, were measured to investigate possible proximate and ultimate mechanisms for the results. The influence of environmental variability was examined, by relating the timing of breeding, survival, and endocrine changes to sea ice conditions. Colony size did not influence the timing or synchrony of breeding, survival, or hormone levels within years; perhaps because colonies in an Adelie rookery are not independent from the 'social environment' of adjacent colonies. Across years, synchrony in the smaller rookery was higher than in the larger rookery. The scale of these comparisons may exceed the applicability of the Darling hypothesis. Therefore, no support was found for the Darling hypothesis, at the colony or rookery level, in this species. Higher corticosterone metabolite and lower sex hormone levels in the first season correlated to later breeding and lower survival compared to the second season. This is likely due to the persistence of extensive sea ice conditions late into the first season.
Researchers should take care in selecting the most appropriate method of hormone measurement for their question. Future studies testing the Darling hypothesis must carefully select their definition of a colony (i.e. a truly isolated social unit) and the scale at which the hypothesis is tested. Combining endocrine measurements with behavioural, survival, and environmental information allows for a more comprehensive interpretation of animal ecology.

year: 2008

title: Testing the boundaries of the middle voice: Observations from English and Romanian

authors: Calude, Andreea S.

subjects: Social Sciences

abstract: The middle voice has received ample attention in the literature, yet the precise boundary between middle voice and other related constructions still remains elusive. For example, do sentences like Mary slept (*herself) well last night and Mary washed (herself) thoroughly and expertly belong to the middle voice or the reflexive domain, or are they simply intransitive one-participant structures? While ambiguity between reflexive and middles has been noted by (Kemmer, Suzanne. 1993. The middle voice. Amsterdam & Philadelphia: John Benjamins.), I show that the problem is more widespread. Depending on the marking patterns available in a given language, such ambiguities can occur between middles and prototypical intransitive one-participant events, or between middles and reflexives. Using data from two languages with distinct marking patterns – English (a language with a reflexive marker but no middle marker) and Romanian (a language with one marker of each type) – I discuss the possibility of distinguishing middles from other related constructions. In English, it is shown that the him/herself test can be used to distinguish direct middles from intransitive one-participant events. In Romanian, we see middle and reflexive markers used together in the same construction without contradiction, bringing together both middle and reflexive semantics (as well as marking). In agreement with (Maldonado, Ricardo. 2000. Spanish reflexives. In Zygmunt Frajzyngier & Traci Walker (eds.), Reflexives: Forms and functions, 153–185. Amsterdam & Philadelphia: John Benjamins.) and (Manney, Linda. 2001. Middle voice in Modern Greek: Meaning and function of a morphoyntactic category. Amsterdam &Philadelphia: John Benjamins.), the account given here supports the view of the middle voice as a unified phenomenon, and following (Maldonado, Ricardo. 2009. Middle as a basic voice system. In Lilian Guerrero, Ibáñez Sergio & Belloro Valeria (eds.), Studies in role and reference grammar. México: Instituto de Investigaciones Filológicas, UNAM.), the main function of the middle is to profile the core properties of events (but not necessarily to reduce the relative elaboration of participants, as proposed by Kemmer, Suzanne. 1993. The middle voice. Amsterdam &Philadelphia: John Benjamins.). So, unlike active and passive voice which concern the focusing of various participants (namely, Agents and Patients, respectively), the middle voice focuses the event itself.

text: Testing the boundaries of the middle voice: Observations from English and Romanian The middle voice has received ample attention in the literature, yet the precise boundary between middle voice and other related constructions still remains elusive. For example, do sentences like Mary slept (*herself) well last night and Mary washed (herself) thoroughly and expertly belong to the middle voice or the reflexive domain, or are they simply intransitive one-participant structures? While ambiguity between reflexive and middles has been noted by (Kemmer, Suzanne. 1993. The middle voice. Amsterdam & Philadelphia: John Benjamins.), I show that the problem is more widespread. Depending on the marking patterns available in a given language, such ambiguities can occur between middles and prototypical intransitive one-participant events, or between middles and reflexives. Using data from two languages with distinct marking patterns – English (a language with a reflexive marker but no middle marker) and Romanian (a language with one marker of each type) – I discuss the possibility of distinguishing middles from other related constructions. In English, it is shown that the him/herself test can be used to distinguish direct middles from intransitive one-participant events. In Romanian, we see middle and reflexive markers used together in the same construction without contradiction, bringing together both middle and reflexive semantics (as well as marking). In agreement with (Maldonado, Ricardo. 2000. Spanish reflexives. In Zygmunt Frajzyngier & Traci Walker (eds.), Reflexives: Forms and functions, 153–185. Amsterdam & Philadelphia: John Benjamins.) and (Manney, Linda. 2001. Middle voice in Modern Greek: Meaning and function of a morphoyntactic category. Amsterdam &Philadelphia: John Benjamins.), the account given here supports the view of the middle voice as a unified phenomenon, and following (Maldonado, Ricardo. 2009. Middle as a basic voice system. In Lilian Guerrero, Ibáñez Sergio & Belloro Valeria (eds.), Studies in role and reference grammar. México: Instituto de Investigaciones Filológicas, UNAM.), the main function of the middle is to profile the core properties of events (but not necessarily to reduce the relative elaboration of participants, as proposed by Kemmer, Suzanne. 1993. The middle voice. Amsterdam &Philadelphia: John Benjamins.). So, unlike active and passive voice which concern the focusing of various participants (namely, Agents and Patients, respectively), the middle voice focuses the event itself.

year: 2017

title: Ngā hau pūkeri e tīhaehae nei i ā tātou tikanga Māori

authors: Nock, Sophie

subjects: Tikanga Māori

abstract: Ko te wherawhera a tēnei tuhinga he titiro ki ngā io mai i ngā tikanga o te ao Māori, ā, ki ngā hau kino e whakataurekareka ana i ā tātou tikanga Māori. Kei roto i ngā iwi katoa o te ao ā rātau tikanga, engari, ko ngā whakamāramatanga o aua tikanga kei te mau ruhiruhi noa i roto i ngā hinengaro o ngā tāngata o tēnā iwi, tēnā iwi, ā, kua ngaro katoa atu rānei. Nā reira, ko tāku he rapu i aua māramatanga kei muri i ngā tikanga Māori, ā, he rapu anō hoki i ngā tāmitanga, i ngā pēhitanga a tauiwi mā, ko te mutunga ka whakapōrearea i ā tātou tikanga Māori.
Ko te aronga rangahau mō tēnei kaupapa he whai i ngā tuhinga me ngā kōrero i ahu mai i te ao Māori me te ao Pākehā. Ko ngā rauemi nei i ahu mai i ngā kohikohinga kōrero tawhito me ngā tuhinga a te hunga o roto i ngā tau kātahi anō ka pahure ake. Ko ngā kōrero tawhito nei hei whakaata ake i te huringa o te ao, ā, ka tīkina atu aua kōrero hei ārahi i te hunga ora kia mau tonu ai ā tātou tikanga Māori.
Ki te whai haere i tēnei momo kaupapa, me mātua whai rawa i ngā kōrero ā-waha hei whakaatu mai, ina kāore hoki i āta tuhia whānuihia ngā kōrero i maringi mai i te hinengaro o te hunga whakautu pātai. Ko ngā kohinga kōrero ā-waha hei tohu i ngā tikanga me ngā whakapono ake i pā mai ki a rātou i te wā e tipu ake ana.
Ko ētahi o ngā kaikōrero i tipu ake i roto i te tūturutanga o te ao Māori, arā, i tipu ake i ngā rekereke o ō rātou kuia, koroua, ā, ko ētahi ahakoa he Māori, i tipu ake anō i roto i te reo me ētahi o ngā tikanga Māori, ko ētahi anō i noho hoki i te tāone.
Ko ngā kaupapa ka whakaarahia ake e tēnei rangahau ko ngā tikanga e hāngai pū ana, e whakamahia ana, e kitea ana, e whakaatu ana anō hoki i ngā whakapono o te Māori. Kāti, ka tirohia ētahi tikanga i waenga i te whānau, ētahi atu tikanga hāngai pū ana ki te manaaki tangata i runga marae me te whakaaro ake ki ngā tohu e tika ana ki te whakanui manuhiri.
He roa te Māori e noho ana i te ao Pākehā, mai i tō rātou taenga mai i te rau tau tekau mā waru. Ko te nuinga o tātau kua roa kē te wehenga mai i te ao tūturu ake o te Māori. Ko ētahi tāngata e pupuri tonu ana i ētahi o ā tātou tikanga, ko ētahi kāhore noa e mōhio ana. Tēnā pea, ko ngā whakatūpato i mau, engari, ko ngā whakamārama kei te ngaro. Kāti, ko te tūmanako mō tēnei tuhingaroa ko te whakahoutia anō o te here ki ā tātou tikanga. Ka āhei hoki te hunga e ngākaunui ana ki te kaupapa nei ki te whai whakaaro, te whai māramatanga.
Ko tēnei tuhinga he tauira hei whakaatu atu he mahi pakeke te hopu, te tiaki, te manaaki me te ako hoki i ngā mātauranga o tuawhakarere. He pēnei anō ēnei tikanga mō nāianei, he toimaha tonu te kawe, te whakaū.
Heoi, ka whakatakia mai ngā kōrero i te orokohanga o te ao hei raranga ake hei whāriki mō te tikanga Māori. Tuatahi, ka tirohia ētahi o ngā whenu o tēnei mahi whata, arā, te ahunga mai o te tikanga Māori me ngā tikanga huhua i whakatauiratia mai e ngā atua Māori. Whai muri atu kei te upoko tuarua, ka whakaraupapahia mai ngā kōrero a te hunga whakautu pātai kia āta kitea ai ō rātou whakaaro mō te tikanga Māori. Ka hāngai pū tonu tēnei titiro ki ngā momo pātai, ā, ka tirohia hoki ngā whakahokinga a taua hunga nā.
Kei te upoko tuatoru ka huri te titiro ki ngā hau pūkeri e tūkino nei i ā tātou tikanga Māori. Tuatahi, ka horahia ngā kōrero, ngā tikanga mō te āhua o te noho a te Māori o mua, kia āta kitea ai i pēhea te āhua o tōna whakahaere i tōna ao o mua. Kātahi, ka āta tirohia ētahi ture Pākehā e tīhaehae nei i ā tātou tikanga, pēnei i te ngaro haere o te reo Māori, te haehae haere o ā tātou tikanga Māori. Ka mutu, ka huri ake ki te whai whakaaro ki ētahi o ngā raru i pā mai ki te tikanga Māori i te neketanga o te Māori ki te tāone noho ai, mahi ai.
Kei te upoko tuawhā ētahi tauira mō ngā tūmomo tikanga e mahia ana e te Māori i waenganui i tōna whānau. Ka tirohia te ringa tapu o te tikanga hei whakaohooho ake i te hinengaro, hei whakatakoto māramatanga mō aua tikanga hoki.
Kei te upoko tuarima te tirohanga ki ngā momo tikanga i runga marae, me ngā momo tohu Māori e whakaatu mai ana i te rangatiratanga o te iwi, pēnei i ngā tohu kino, i ngā tohu ora rānei. Ka tirohia ētahi whakamāramatanga mō te āhua o te noho piri o te Māori ki tōna taiao, he aha ngā kōrero a te taiao ki te Māori, he aha hoki ngā kōrero a ngā kararehe o te taiao.
Hei whakakapi ake i te tuhinga nei ko te whakarāpopototanga. I konei ka titiro whakamuri ki ngā āhuatanga kua ara ake. Ka whakaraupapahia ngā whakakitenga me ngā whakaputanga hei whakaohooho, hei whakaarotanga ake mō te ao kei mua e pā ana ki ā tātou tikanga Māori.

text: Ngā hau pūkeri e tīhaehae nei i ā tātou tikanga Māori Ko te wherawhera a tēnei tuhinga he titiro ki ngā io mai i ngā tikanga o te ao Māori, ā, ki ngā hau kino e whakataurekareka ana i ā tātou tikanga Māori. Kei roto i ngā iwi katoa o te ao ā rātau tikanga, engari, ko ngā whakamāramatanga o aua tikanga kei te mau ruhiruhi noa i roto i ngā hinengaro o ngā tāngata o tēnā iwi, tēnā iwi, ā, kua ngaro katoa atu rānei. Nā reira, ko tāku he rapu i aua māramatanga kei muri i ngā tikanga Māori, ā, he rapu anō hoki i ngā tāmitanga, i ngā pēhitanga a tauiwi mā, ko te mutunga ka whakapōrearea i ā tātou tikanga Māori.
Ko te aronga rangahau mō tēnei kaupapa he whai i ngā tuhinga me ngā kōrero i ahu mai i te ao Māori me te ao Pākehā. Ko ngā rauemi nei i ahu mai i ngā kohikohinga kōrero tawhito me ngā tuhinga a te hunga o roto i ngā tau kātahi anō ka pahure ake. Ko ngā kōrero tawhito nei hei whakaata ake i te huringa o te ao, ā, ka tīkina atu aua kōrero hei ārahi i te hunga ora kia mau tonu ai ā tātou tikanga Māori.
Ki te whai haere i tēnei momo kaupapa, me mātua whai rawa i ngā kōrero ā-waha hei whakaatu mai, ina kāore hoki i āta tuhia whānuihia ngā kōrero i maringi mai i te hinengaro o te hunga whakautu pātai. Ko ngā kohinga kōrero ā-waha hei tohu i ngā tikanga me ngā whakapono ake i pā mai ki a rātou i te wā e tipu ake ana.
Ko ētahi o ngā kaikōrero i tipu ake i roto i te tūturutanga o te ao Māori, arā, i tipu ake i ngā rekereke o ō rātou kuia, koroua, ā, ko ētahi ahakoa he Māori, i tipu ake anō i roto i te reo me ētahi o ngā tikanga Māori, ko ētahi anō i noho hoki i te tāone.
Ko ngā kaupapa ka whakaarahia ake e tēnei rangahau ko ngā tikanga e hāngai pū ana, e whakamahia ana, e kitea ana, e whakaatu ana anō hoki i ngā whakapono o te Māori. Kāti, ka tirohia ētahi tikanga i waenga i te whānau, ētahi atu tikanga hāngai pū ana ki te manaaki tangata i runga marae me te whakaaro ake ki ngā tohu e tika ana ki te whakanui manuhiri.
He roa te Māori e noho ana i te ao Pākehā, mai i tō rātou taenga mai i te rau tau tekau mā waru. Ko te nuinga o tātau kua roa kē te wehenga mai i te ao tūturu ake o te Māori. Ko ētahi tāngata e pupuri tonu ana i ētahi o ā tātou tikanga, ko ētahi kāhore noa e mōhio ana. Tēnā pea, ko ngā whakatūpato i mau, engari, ko ngā whakamārama kei te ngaro. Kāti, ko te tūmanako mō tēnei tuhingaroa ko te whakahoutia anō o te here ki ā tātou tikanga. Ka āhei hoki te hunga e ngākaunui ana ki te kaupapa nei ki te whai whakaaro, te whai māramatanga.
Ko tēnei tuhinga he tauira hei whakaatu atu he mahi pakeke te hopu, te tiaki, te manaaki me te ako hoki i ngā mātauranga o tuawhakarere. He pēnei anō ēnei tikanga mō nāianei, he toimaha tonu te kawe, te whakaū.
Heoi, ka whakatakia mai ngā kōrero i te orokohanga o te ao hei raranga ake hei whāriki mō te tikanga Māori. Tuatahi, ka tirohia ētahi o ngā whenu o tēnei mahi whata, arā, te ahunga mai o te tikanga Māori me ngā tikanga huhua i whakatauiratia mai e ngā atua Māori. Whai muri atu kei te upoko tuarua, ka whakaraupapahia mai ngā kōrero a te hunga whakautu pātai kia āta kitea ai ō rātou whakaaro mō te tikanga Māori. Ka hāngai pū tonu tēnei titiro ki ngā momo pātai, ā, ka tirohia hoki ngā whakahokinga a taua hunga nā.
Kei te upoko tuatoru ka huri te titiro ki ngā hau pūkeri e tūkino nei i ā tātou tikanga Māori. Tuatahi, ka horahia ngā kōrero, ngā tikanga mō te āhua o te noho a te Māori o mua, kia āta kitea ai i pēhea te āhua o tōna whakahaere i tōna ao o mua. Kātahi, ka āta tirohia ētahi ture Pākehā e tīhaehae nei i ā tātou tikanga, pēnei i te ngaro haere o te reo Māori, te haehae haere o ā tātou tikanga Māori. Ka mutu, ka huri ake ki te whai whakaaro ki ētahi o ngā raru i pā mai ki te tikanga Māori i te neketanga o te Māori ki te tāone noho ai, mahi ai.
Kei te upoko tuawhā ētahi tauira mō ngā tūmomo tikanga e mahia ana e te Māori i waenganui i tōna whānau. Ka tirohia te ringa tapu o te tikanga hei whakaohooho ake i te hinengaro, hei whakatakoto māramatanga mō aua tikanga hoki.
Kei te upoko tuarima te tirohanga ki ngā momo tikanga i runga marae, me ngā momo tohu Māori e whakaatu mai ana i te rangatiratanga o te iwi, pēnei i ngā tohu kino, i ngā tohu ora rānei. Ka tirohia ētahi whakamāramatanga mō te āhua o te noho piri o te Māori ki tōna taiao, he aha ngā kōrero a te taiao ki te Māori, he aha hoki ngā kōrero a ngā kararehe o te taiao.
Hei whakakapi ake i te tuhinga nei ko te whakarāpopototanga. I konei ka titiro whakamuri ki ngā āhuatanga kua ara ake. Ka whakaraupapahia ngā whakakitenga me ngā whakaputanga hei whakaohooho, hei whakaarotanga ake mō te ao kei mua e pā ana ki ā tātou tikanga Māori.

year: 2003