Placental morphology in two sympatric Andean lizards of the genus Liolaemus (Reptilia: Liolaemidae)

Viviparity is a remarkable feature in squamate sauropsids and it has evolved multiple times in parallel with the formation of a placenta. One example of this repeated evolution of viviparity and placentation occurs in the species‐rich South American genus Liolaemus with at least six independent origins of viviparity. However, evolutionary studies of placentation in this genus are limited by a lack of data on placental morphology. The aim of this study is to describe and compare the microanatomy and vessel diameter (Dv, a function of blood flow) of the placenta using scanning electron microscopy (SEM) and confocal laser scanning microscopy (cLSM) in two sympatric Andean viviparous but highly divergent species, Liolaemus robustus and Liolaemus walkeri. We found interspecific differences in cell types in the chorion, allantois, and omphalopleure that may be explained by divergent phylogenetic history. Time elapsed since divergence may also explain the pronounced interspecific differences in vessel diameter, and within each species, there are strong differences in Dv between tissue locations. Both species show features to improve gas exchange in the chorioallantoic placenta including absence of eggshell, large Dv in the allantois (L. robustus) or embryonic side of the uterus (L. walkeri), and when present, microvillous cells in the allantois (L. walkeri). Both species also show features that suggest transfer of nutrients or water in the omphaloplacenta, including an almost complete reduction of the eggshell, secretive material (L. robustus), or vesicles (L. walkeri) on cell surface uterus, and when present specialized cells in the omphalopleure (L. walkeri). No statistical differences in Dv were found among stages 32–39 in each species, suggesting that a different mechanism, other than enhanced blood flow, might satisfy the increased oxygen demand of the developing embryos in the hypoxic environments of the high Andes. J. Morphol. 276:1205–1217, 2015. © 2015 Wiley Periodicals, Inc.     

Thermal biology and locomotor performance of the Andean lizard Liolaemus fitzgeraldi (Liolaemidae) in Argentina

Ectotherms thermoregulate to maintain their body temperature within the optimal range needed for performing vital functions. The effect of climate change on lizards has been studied as regards the sensitivity of locomotor performance to environmental temperatures. We studied thermoregulatory efficiency and locomotor performance for Liolaemus fitzgeraldi in the Central Andes of Argentina. We determined body temperature, micro-environmental temperatures and operative temperatures in the field. In the laboratory, we measured preferred temperatures and calculated the index of thermoregulatory efficiency. We estimated the thermal sensitivity of locomotion by measuring sprint speed (initial velocity and long sprint) and endurance at five different body temperatures. Body temperature was not associated with either micro-environmental temperature, nor did it show differences with preferred temperatures. Thermoregulatory efficiency was moderate (0.61). Initial velocity and long sprint trials showed differences at different temperatures; however, endurance did not. Moreover, the optimal temperatures for the performance trials showed no significant differences among themselves. We conclude that Liolaemus fitzgeraldi has thermal sensitivity in locomotor performance with respect to body temperature and that it is an eurythermic lizard that experiences a large variation in body temperature and that has thermal flexibility in the cold.     

Population Dynamics Following the Last Glacial Maximum in Two Sympatric Lizards in Northern China

Phylogeographic studies of Eremias lizards （Lacertidae） in East Asia have been limited, and the impact of major climatic events on their population dynamics remains poorly known. This study aimed to investigate population histories and refugia during the Last Glacial Maximum of two sympatric Eremias lizards （E. argus and E. brenchleyi） inhabiting northern China. We sequenced partial mitochondrial DNA from the ND4 gene for 128 individuals of E. argus from nine localities, and 46 individuals of E. brenchleyi from five localities. Forty-four ND4 haplotypes were determined from E. argus samples, and 33 from E. brenchleyi samples. Population expansion events began about 0.0044 Ma in E. argus, and 0.031 Ma in E. brenchleyi. The demographic history of E. brenchleyi indicates a long-lasting population decline since the most recent common ancestor, while that of E. argus indicates a continuous population growth. Among-population structure was significant in both species, and there were multiple refugia across their range. Intermittent gene flow occurred among expanded populations across multiple refugia during warmer phases of the glacial period, and this may explain why the effective population size has remained relatively stable in E. brenchleyi and grown in E. argus.     

末次盛冰期环境恶化对中国北方旧石器文化的影响

末次冰期间冰段时,中国北方地区水热条件普遍较好,旧石器文化也比较发达。然而,进入末次盛冰期后,随着气温的普遍降低和降水量的明显减少,草原和沙漠带急速向南扩张,动物群也大举南迁。对于这种严酷的生存环境,中国北方的旧石器文化是如何响应的呢?本文以宁夏彭阳PY01-02,PY03和PY05等地点的地层剖面和石制品的研究为基础,首先提出彭阳地点群地层沉积相由河湖相沉积迅速转变为黄土堆积说明由末次冰期间冰段到末次盛冰期自然环境发生了剧烈变化。其次,通过对上述两种不同沉积相中出土石制品的大小、重量、原料以及技术上的比较说明随着环境的变化石制品本身也发生了相应的变化。下层河湖相堆积中出土的石制品粗大笨重,主要以各种石英岩为原料,与同期泾水上游旧石器文化的特征相近,应是当地文化特征的一种具体体现。上层马兰黄土中的石制品的普遍较小,主要以石英和燧石为原料,其中用燧石制成的细石器具有宁夏灵武水洞沟下文化层同类制品的特征,而用石英制成的小石器则同海原一带的史前文化有着较为密切的联系,据此作者推测这可能与史前人类在环境日益恶劣的情况下向南迁移有关。山西南部地区的下川和柿子滩遗址的地层和石制品证据亦表明,末次盛冰期前后该地区旧石器文化的明显差异可能也与末次盛冰期环境恶化时人类的迁徙有关。最后,中国北方地区考古遗址的空间分布变化为上述史前人类南迁的推测提供了有力证据。因此,作者认为末次冰期的环境恶化对中国北方旧石器文化的影响具有普遍性。     

Evidence for a lacustrine faunal refuge in the Larsemann Hills, East Antarctica, during the Last Glacial Maximum

Aim  There is no previous direct evidence for the occurrence of lacustrine refuges for invertebrate fauna in Antarctica spanning the Last Glacial Maximum (LGM). In the absence of verified LGM lacustrine refuges many species are believed to result from Holocene dispersal from sub-Antarctic islands and continents further north. If freshwater lake environments were present throughout the LGM, extant freshwater species may have been associated with Antarctica prior to this glacial period. This study looked at faunal microfossils in a sediment core from an Antarctic freshwater lake. This lake is unusual in that, unlike most Antarctic lakes, the sediment record extends to c . 130,000 yr bp , i.e. prior to the LGM.
Location  Lake Reid, Larsemann Hills, East Antarctica (76°23' E; 69°23' S).
Methods  Palaeofaunal communities in Lake Reid were identified through examination of faunal microfossils in a sediment core that extended to c . 130,000 yr bp .
Results  Ephippia and mandibles from the cladoceran Daphniopsis studeri and loricae of the rotifer Notholca sp. were found at all depths in the sediment, indicating that these two species have been present in the lake for up to 130,000 years. Copepod mandibles were also present in the older section of the core, yet were absent from the most recent sediments, indicating extinction of this species from Lake Reid during the LGM.
Main conclusion  The presence of D. studeri and Notholca sp. microfossils throughout the entire Lake Reid core is the first direct evidence of a glacial lacustrine refugium for invertebrate animals in Antarctica, and indicates the presence of a relict fauna on the Antarctic continent.     

Satsurblia: New Insights of Human Response and Survival across the Last Glacial Maximum in the Southern Caucasus

The region of western Georgia (Imereti) has been a major geographic corridor for human migrations during the Middle and Upper Palaeolithic (MP/UP). Knowledge of the MP and UP in this region, however, stems mostly from a small number of recent excavations at the sites of Ortvale Klde, Dzudzuana, Bondi, and Kotias Klde. These provide an absolute chronology for the Late MP and MP–UP transition, but only a partial perspective on the nature and timing of UP occupations, and limited data on how human groups in this region responded to the harsh climatic oscillations between 37,000–11,500 years before present. Here we report new UP archaeological sequences from fieldwork in Satsurblia cavein the same region. A series of living surfaces with combustion features, faunal remains, stone and bone tools, and ornaments provide new information about human occupations in this region (a) prior to the Last Glacial Maximum (LGM) at 25.5–24.4 ka cal. BP and (b) after the LGM at 17.9–16.2 ka cal. BP. The latter provides new evidence in the southern Caucasus for human occupation immediately after the LGM. The results of the campaigns in Satsurblia and Dzudzuana suggest that at present the most plausible scenario is one of a hiatus in the occupation of this region during the LGM (between 24.4–17.9 ka cal. BP). Analysis of the living surfaces at Satsurblia offers information about human activities such as the production and utilisation of lithics and bone tools, butchering, cooking and consumption of meat and wild cereals, the utilisation of fibers, and the use of certain woods. Microfaunal and palynological analyses point to fluctuations in the climate with consequent shifts in vegetation and the faunal spectrum not only before and after the LGM, but also during the two millennia following the end of the LGM.     

Study of lipid reserves in Liolaemus koslowskyi (Squamata: Liolaemidae): reproductive and ecological implications

Journal of Comparative Physiology B - In reptiles, many lipid reserve structures were recognized, and different patterns of storage and utilization of lipids have been identified. In this study, a...     

Phylogeny and phylogeography of the Liolaemus darwinii complex (Squamata: Liolaemidae): evidence for introgression and incomplete lineage sorting

Although mitochondrial DNA markers have several properties that make them suitable for phylogeographic studies, they are not free of difficulties. Phylogeographic inferences within and between closely related species can be mislead by introgression and retention of ancestral polymorphism. Here we combine different phylogenetic, phylogeographic, and population genetic methods to extract the maximum information from the Liolaemus darwinii complex. We estimate the phylogeographic structure of L. darwinii across most of its distributional range, and we then estimate relationships between L. darwinii and the syntopic species L. laurenti and L. grosseorum. Our results suggest that range expansion of these lineages brought them into secondary contact in areas where they are presently in syntopy. Here we present the first evidence for introgression in lizards from temperate South America (of L. danwinii mitochondrial DNA into L. laurenti and L. grosseorum), and for incomplete lineage sorting (between L. darwinii and L. laurenti). We show that a combination of methods can provide additional support for inferences derived from any single method and thus provide more robust interpretations and narrow the range of plausible hypotheses about mechanisms and processes of divergence. Additional studies are needed in this group of lizards and in other codistributed groups to determine if Pleistocene climatic changes could be a general factor influencing the evolutionary history of a regional biota.     

A new species of Liolaemus related to L. nigroviridis from the Andean highlands of Central Chile (Iguania,Liolaemidae)

The Liolaemus nigroviridis group is a clade of highland lizards endemic to Chile. These species are distributed from northern to central Chile, and currently there are no cases of sympatric distribution. This study describes a new species, Liolaemus uniformis sp. n., from this group, and provides a detailed morphological characterization and mitochondrial phylogeny using cytochrome-b. Liolaemus uniformis was found in sympatry with Liolaemus nigroviridis but noticeably differed in size, scalation, and markedly in the color pattern, without sexual dichromatism. This new species has probably been confused with Liolaemus monticola and Liolaemus bellii, both of which do not belong to the nigroviridis group. The taxonomic issues of this group that remain uncertain are also discussed.     

Genetic and palaeo-climatic evidence for widespread persistence of the coastal tree species Eucalyptus gomphocephala (Myrtaceae) during the Last Glacial Maximum

Background and Aims

Few phylogeographic studies have been undertaken of species confined to narrow, linear coastal systems where past sea level and geomorphological changes may have had a profound effect on species population sizes and distributions. In this study, a phylogeographic analysis was conducted of Eucalyptus gomphocephala (tuart), a tree species restricted to a 400 × 10 km band of coastal sand-plain in south west Australia. Here, there is little known about the response of coastal vegetation to glacial/interglacial climate change, and a test was made as to whether this species was likely to have persisted widely through the Last Glacial Maximum (LGM), or conforms to a post-LGM dispersal model of recovery from few refugia.

Methods

The genetic structure over the entire range of tuart was assessed using seven nuclear (21 populations; n = 595) and four chloroplast (24 populations; n = 238) microsatellite markers designed for eucalypt species. Correlative palaeodistribution modelling was also conducted based on five climatic variables, within two LGM models.

Key Results

The chloroplast markers generated six haplotypes, which were strongly geographically structured (G_ST = 0·86 and R_ST = 0·75). Nuclear microsatellite diversity was high (overall mean H_E 0·75) and uniformly distributed (F_ST = 0·05), with a strong pattern of isolation by distance (r² = 0·362, P = 0·001). Distribution models of E. gomphocephala during the LGM showed a wide distribution that extended at least 30 km westward from the current distribution to the palaeo-coastline.

Conclusions

The chloroplast and nuclear data suggest wide persistence of E. gomphocephala during the LGM. Palaeodistribution modelling supports the conclusions drawn from genetic data and indicates a widespread westward shift of E. gomphocephala onto the exposed continental shelf during the LGM. This study highlights the importance of the inclusion of complementary, non-genetic data (information on geomorphology and palaeoclimate) to interpret phylogeographic patterns.     

Microhabitat use,daily activity pattern,and diet of Liolaemus etheridgei Laurent, 1998 (Reptilia: Liolaemidae) in the Andean Polylepis forests of Arequipa,Peru

This study describes the microhabitat use, daily activity pattern, and diet of Liolaemus etheridgei Laurent, 1998 in the El Simbral and Tuctumpaya Polylepis forests in Arequipa, Peru. El Simbral is a fragmented forest, whereas Tuctumpaya is unfragmented. Our results reveal that L. etheridgei shows no positive selection for any of the microhabitats we identified in Polylepis forests; on the contrary, it selects negatively against Polylepis trees and nonthorny bushes. The daily activity patterns indicate a bimodal pattern with peaks at 9:00–10:59 and 13:00–13:59 h. The diet of L. etheridgei consists mainly of plant material, and the most important animal prey category is Lygaeidae: Hemiptera, which is selected for positively. In particular, microhabitat selection varied for nonthorny bushes, which were selected negatively in the Tuctumpaya population but neither positively nor negatively in the El Simbral population. According to the proportions of plant material found, the L. etheridgei from El Simbral were found to be omnivorous, whereas the Tuctumpaya population was herbivorous. However, the percentage of plant material consumed in the El Simbral population was close to the critical value for herbivory–omnivory. We conclude that the three ecological aspects of L. etheridgei studied here are virtually identical in El Simbral and Tuctumpaya; therefore, this species is not affected significantly by the current fragmentation of forest.     

Predictive modelling of tree species distributions on the Iberian Peninsula during the Last Glacial Maximum and Mid-Holocene

This paper reports a bioclimatic envelope model study of the potential distribution of 19 tree species in the Iberian Peninsula during the Last Glacial Maximum (LGM; 21 000 yr BP) and the Mid-Holocene (6000 yr BP). Current patterns of tree species richness and distributions are believed to have been strongly influenced by the climate during these periods. The modelling employed novel machine learning techniques, and its accuracy was evaluated using a threshold-independent method. Two atmospheric general circulation models, UGAMP and ECHAM3 (generated by the Palaeoclimate Modelling Intercomparison Project, PMIP), were used to provide climate scenarios under which the distributions of the 19 tree species were modelled. The results obtained for these scenarios were assessed by agreement measure analysis; they differed significantly for the LGM, but were more similar for the Mid-Holocene.
The results for the LGM support the inferred importance of pines in the Iberian Peninsula at this time, and the presence of evergreen Quercus in the south. Important differences in the altitude at which the modelled species grew were also predicted. During the LGM, some normally higher mountain species potentially became re-established in the foothills of the Pyrenees. The warm Mid-Holocene climate is clearly reflected in the predicted expansion of broad-leaved forests during this period, including the colonization of the northern part of the Iberian Peninsula by evergreen Quercus species.     

Sex‐specific evolution of bite performance in Liolaemus lizards (Iguania: Liolaemidae): the battle of the sexes

Although differential selective pressures on males and females of the same species may result in sex‐specific evolutionary trajectories, comparative studies of adaptive radiations have largely neglected within‐species variation. In this study, we explore the potential effects of natural selection, sexual selection, or a combination of both, on bite performance in males and females of 19 species of Liolaemus lizards. More specifically, we study the evolution of bite performance, and compare evolutionary relationships between the variation in head morphology, bite performance, ecological variation and sexual dimorphism between males and females. Our results suggest that in male Liolaemus, the variation in bite force is at least partly explained by the variation in the degree of sexual dimorphism in head width (i.e. our estimate of the intensity of sexual selection), and neither bite force nor the morphological variables were correlated with diet (i.e. our proxy for natural selection). On the contrary, in females, the variation in bite force and head size can, to a certain extent, be explained by variation in diet. These results suggest that whereas in males, sexual selection seems to be operating on bite performance, in the case of females, natural selection seems to be the most likely and most important selective pressure driving the variation in head size. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 101 , 461–475.     

Relaxed Molecular Clock Provides Evidence for Long-Distance Dispersal of Nothofagus (Southern Beech)

Nothofagus (southern beech), with an 80-million-year-old fossil record, has become iconic as a plant genus whose ancient Gondwanan relationships reach back into the Cretaceous era. Closely associated with Wegener's theory of “Kontinentaldrift”, Nothofagus has been regarded as the “key genus in plant biogeography”. This paradigm has the New Zealand species as passengers on a Moa's Ark that rafted away from other landmasses following the breakup of Gondwana. An alternative explanation for the current transoceanic distribution of species seems almost inconceivable given that Nothofagus seeds are generally thought to be poorly suited for dispersal across large distances or oceans. Here we test the Moa's Ark hypothesis using relaxed molecular clock methods in the analysis of a 7.2-kb fragment of the chloroplast genome. Our analyses provide the first unequivocal molecular clock evidence that, whilst some Nothofagus transoceanic distributions are consistent with vicariance, trans-Tasman Sea distributions can only be explained by long-distance dispersal. Thus, our analyses support the interpretation of an absence of Lophozonia and Fuscospora pollen types in the New Zealand Cretaceous fossil record as evidence for Tertiary dispersals of Nothofagus to New Zealand. Our findings contradict those from recent cladistic analyses of biogeographic data that have concluded transoceanic Nothofagus distributions can only be explained by vicariance events and subsequent extinction. They indicate that the biogeographic history of Nothofagus is more complex than envisaged under opposing polarised views expressed in the ongoing controversy over the relevance of dispersal and vicariance for explaining plant biodiversity. They provide motivation and justification for developing more complex hypotheses that seek to explain the origins of Southern Hemisphere biota.     

Phylogeographical Analysis of mtDNA Data Indicates Postglacial Expansion from Multiple Glacial Refugia in Woodland Caribou (Rangifer tarandus caribou)

Glacial refugia considerably shaped the phylogeographical structure of species and may influence intra-specific morphological, genetic, and adaptive differentiation. However, the impact of the Quaternary ice ages on the phylogeographical structure of North American temperate mammalian species is not well-studied. Here, we surveyed ∼1600 individuals of the widely distributed woodland caribou (Rangifer tarandus caribou) using mtDNA control region sequences to investigate if glacial refugia contributed to the phylogeographical structure in this subspecies. Phylogenetic tree reconstruction, a median-joining network, and mismatch distributions supported postglacial expansions of woodland caribou from three glacial refugia dating back to 13544–22005 years. These three lineages consisted almost exclusively of woodland caribou mtDNA haplotypes, indicating that phylogeographical structure was mainly shaped by postglacial expansions. The putative centres of these lineages are geographically separated; indicating disconnected glacial refugia in the Rocky Mountains, east of the Mississippi, and the Appalachian Mountains. This is in congruence with the fossil record that caribou were distributed in these areas during the Pleistocene. Our results suggest that the last glacial maximum substantially shaped the phylogeographical structure of this large mammalian North American species that will be affected by climatic change. Therefore, the presented results will be essential for future conservation planning in woodland caribou.     

Phylogeographic analyses suggest that a deciduous species (Ostryopsis davidiana Decne., Betulaceae) survived in northern China during the Last Glacial Maximum

Aim Palaeontologial data suggest that all temperate forest species in northern China migrated southwards during the Last Glacial Maximum (LGM) and recolonized post‐glacially within the Holocene. We tested this assumption using phylogeographical studies of a temperate deciduous shrub species (Ostryopsis davidiana Decne., Betulaceae), which has a wide distribution in northern China. Location Northern China. Methods We sequenced two chloroplast DNA (cpDNA) fragments (trnL–trnF and psbA–trnH, together about 1300 bp in length) of 294 plants from 21 populations across the total distribution range of this species. We used maximum parsimony and haplotype network methods to construct phylogenetic relationships among haplotypes. Results The analysis of cpDNA variation identified eight haplotypes. A single haplotype was fixed in all populations except for one population that was polymorphic, having two haplotypes. The population subdivisions were extremely high (G_ST = 0.972 and N_ST = 0.974), suggesting very low gene flow between populations. Haplotypes clustered into two tentative clades, both of which occur in the southern region of the species’ range but only one of which occurs in the northern region. Across the sampled populations, the haplotype distributions were differentiated geographically. Main conclusions Our analyses suggest that multiple refugia were maintained across the range of O. davidiana in both northern (north of the Qing Mountains) and southern (south of the Qing Mountains) regions during the LGM rather than that the species survived only in the south and subsequently colonized northwards. The extremely low within‐population diversity of this species suggests strong bottleneck or founder effects within each fragmented region during the Quaternary climatic oscillations. These findings provide important clues for understanding range shifts and changes in within‐ and/or between‐population genetic diversity of temperate forests in response to past climatic oscillations in northern China.