Brief communication: Forelimb compliance in arboreal and terrestrial opossums

Primates display high forelimb compliance (increased elbow joint yield) compared to most other mammals. Forelimb compliance, which is especially marked among arboreal primates, moderates vertical oscillations of the body and peak vertical forces and may represent a basal adaptation of primates for locomotion on thin, flexible branches. However, Larney and Larson (Am J Phys Anthropol 125 [2004] 42–50) reported that marsupials have forelimb compliance comparable to or greater than that of most primates, but did not distinguish between arboreal and terrestrial marsupials. If forelimb compliance is functionally linked to locomotion on thin branches, then elbow yield should be highest in marsupials relying on arboreal substrates more often. To test this hypothesis, we compared forelimb compliance between two didelphid marsupials, Caluromys philander (an arboreal opossum relying heavily on thin branches) and Monodelphis domestica (an opossum that spends most of its time on the ground). Animals were videorecorded while walking on a runway or a horizontal 7‐mm pole. Caluromys showed higher elbow yield (greater changes in degrees of elbow flexion) on both substrates, similar to that reported for arboreal primates. Monodelphis was characterized by lower elbow yield that was intermediate between the values reported by Larney and Larson (Am J Phys Anthropol 125 [2004] 42–50) for more terrestrial primates and rodents. This finding adds evidence to a model suggesting a functional link between arboreality—particularly locomotion on thin, flexible branches—and forelimb compliance. These data add another convergent trait between arboreal primates, Caluromys, and other arboreal marsupials and support the argument that all primates evolved from a common ancestor that was a fine‐branch arborealist. Am J Phys Anthropol, 2010. © 2009 Wiley‐Liss, Inc.     

Stride lengths and frequencies of arboreal walking in seven species of didelphid marsupials

Didelphid marsupials differ in their use of the forest strata, with corresponding differences in morphology and arboreal walking performances. Similar performances may be reached by different combinations of stride length and frequency, but it has been suggested that arboreal walkers increase velocity by longer strides. Our objective was to determine how stride length and frequency contribute to the velocity in the arboreal walking of seven species of didelphid marsupials of the Atlantic Forest of Brazil. Animals were stimulated to cross five 3-m long horizontal supports of different diameters. The cycle of maximum velocity was chosen to measure relative stride length, frequency, and relative velocity. Except forCaluromys philander, the more arboreal species were faster than the terrestrial species, but maximum velocity of arboreal species was reached by two strategies, increasing stride frequency (Gracilinanus microtarsus, Micoureus demerarae, andDidelphis aurita), or reducing frequency and increasing stride length (Marmosops incanus andC. philander). Increasing velocity in arboreal walking by more frequent strides may reduce oscillations of the body, whereas longer strides may reduce branch swaying. Among the terrestrial species,Philander frenatus performed similarly to more arboreal species, suggesting a potential ability to use the canopy, undetected in field observations.     

Use of zygodactylous grasp by Caluromys philander (Didelphimorphia: Didelphidae)

The present report aims to quantify the use of zygodactylous (opposability of digits II to III) grasping in relation to positional modes and support size and orientation, in the highly arboreal, walking/climbing woolly opossum, Caluromys philander. For this purpose, four captive adult C. philander were intensively video-recorded and their positional behavior, hand grasp, and support size and orientation use were analyzed frame-by-frame. Overall, C. philander used a zygodactylous grasp in 81.3±1.2% of bouts. In terms of support features, this grasp was particularly common on (a) supports that could be wholly and partly held by the animals’ hand and (b) vertical supports in particular. In a comparable manner, zygodactyly dominated during above-support positional modes, but was significantly less used during bridging and suspension. The results show that zygodactyly provided an above-support secure and steady grasp on relatively unstable arboreal supports, by aligning the hand with the main axis of the support. This very likely assisted in controlling over the applied torques during cautious quadrupedal and climbing activities with extended hand contact that characterizes the locomotor strategy of C. philander. These observations need to be further tested by more detailed kinetic studies and on a larger sample of arboreal didelphids.     

Use of forest canopy by European red squirrelsSciurus vulgaris in Northern Greece: claws and the small branch niche

Moving and standing in trees impose multiple problems to arboreal mammals. Among them, the major ones are the negotiation of slender terminal branches and of large vertical supports. Both microhabitats are important as they have been linked alternatively to the evolutionary loss of claws in early primates. Therefore, rates of use of these different supports by claw-bearing arboreal mammals may offer insights to their actual significance in the adaptive evolution of early primates. In this context, canopy, tree crown, branch size, inclination, and texture use were recorded on four adult free ranging European red squirrelsSciurus vulgaris Linnaeus, 1758 in a mixed coniferous forest in northern Greece.S. vulgaris was mainly arboreal, exploiting the terminal branch zone, using frequently oblique and intermediately textured supports<5 cm and moderately large vertical branches. Furthermore, comparative data from other sciurid species and clawed primates showed positive correlations of small and horizontal support use, and negative ones of vertical support use to body mass. These findings show that keeled functional claws do not impede habitual use of slender branches and may not facilitate efficient climbing on large vertical trunks. These observations partly question the association between habitual use of the small branch niche and primate adaptations and lend support to alternative hypotheses, underscoring the importance of inquiring for more complex mechanisms that lead to the evolution of the unique set of primate morphological adaptations.     

Arboreal walking performance in seven didelphid marsupials as an aspect of their fundamental niche

Abstract Species of didelphid marsupials (Didelphimorphia, Didelphidae) differ in their use of the forest strata, but it is not clear whether these differences are in fundamental or realized niches. The fundamental niche of seven species of didelphids (Caluromys philander, Didelphis aurita, Gracilinanus microtarsus, Marmosops incanus, Metachirus nudicaudatus, Micoureus demerarae, and Philander frenatus) was compared using their performance in arboreal walking. The association between performance and vertical use of the forest also was tested accounting for phylogenetic and allometric effects. Tests consisted of making the animal cross five 3 m long horizontal supports of different diameters, 1 m from the ground. The cycle of maximum speed was chosen to measure stride length, frequency and velocity. Arboreal species performed better than the terrestrial ones, but a major part of the variation in stride length (70.95%) and stride frequency (88.10%) was associated with body size. Part of the variation in stride length independent of body size (14.05%) was associated with the degree of vertical use of the forest, after phylogenetic effects were accounted for. Fundamental niches of six of the seven species were discriminated with the performance tests used. Discrepancies between the realized and fundamental niches can be inferred for two of these species, D. aurita and P. frenatus.     

Locomotion and Posture During Terminal Branch Feeding

We investigated locomotor and postural behavior during terminal branch feeding in order to gain a better understanding of the motor capabilities of primates. We videotaped wild, juvenile bonnet macaques (Macaca radiata) in India as they fed on flower nectar in a simal tree (Bombax malabaricum). Kinematic analysis revealed that they select specific support surfaces and movements that, for their body design and postures, maximize lateral stability and minimize the chances of falling. These choices are made even though the distance and duration of travel to a selected target are frequently increased. Our discussion focuses on particular concepts of how primates contend with balance problems arboreally, potential reasons for changes in footfall patterns, and how the tail contributes to arboreal locomotion and posture. We concluded that balance problems due to the ratio of body size to branch size are usually avoided, at least among juvenile bonnet macaques, by placing the hands and feet on branches extending laterally from the central support branch and not on the central branch itself. The lateral branches permit a wide base of support, which increases lateral stability. Second, juvenile bonnet macaques have a striking ability to rapidly and repeatedly adapt their gait patterns to changing substrate design with minimal interruption to overall progression. Third, primate tails that are not morphologically specialized for prehension nevertheless have important prehensile and sensory functions in arboreal locomotion and posture.     

Morphometrics confirm taxonomic deflation in dwarf lemurs (Primates: Cheirogaleidae), as suggested by genetics

Detailed knowledge of the identity and distributions of extant species is essential for unravelling patterns and mechanisms of biodiversity, and it provides indispensable baseline data for conservation efforts. The taxonomy of the primates of Madagascar (Lemuriformes) has experienced drastic revisions in the last decades, with species numbers skyrocketing, especially in the genera Microcebus, Avahi, and Lepilemur. The dwarf lemurs (genus Cheirogaleus) have received less attention in terms of their taxonomy, even though they are closely related and syntopic with these diverse genera. The last revision of the genus was based on morphological data from museum specimens, and accepted seven species: Cheirogaleus medius, Cheirogaleus major, Cheirogaleus crossleyi, Cheirogaleus adipicaudatus, Cheirogaleus sibreei, Cheirogaleus ravus, and Cheirogaleus minusculus, whereas a more recent multilocus molecular study only detected three lineages: C. medius, C. major, and C. crossleyi. The goal of this study was to reassess the currently accepted taxonomy of the genus by examining six external and 32 craniodental characters of 120 museum specimens and 36 new specimens from the field. This study, in conjunction with a complementary molecular study, revealed lower diversity and a lower number of distinct morphs of dwarf lemurs than previously postulated. We conclude that in our sample there are three distinct morphs in the genus Cheirogaleus that correspond to C. medius, C. major, and C. crossleyi. We formally recognize C. adipicaudatus and C. ravus as synonyms of C. medius and C. major, respectively, and consider C. minusculus and C. sibreei to be potential synonyms of C. medius. © 2010 The Linnean Society of London, Zoological Journal of the Linnean Society, 2010.     

Primate origins: Lessons from a neotropical marsupial

The didelphid Caluromys shows evolutionary convergence towards prosimians in having a relatively large brain, large eyes, small litters, slow development, and agile locomotion. The selection pressures that favored the emergence of primate-like traits in Caluromys from a generalized didelphid ancestor may be analogous to the selection pressures favoring the initial divergence of primates from a primitive nonprimate ancestor, and thus Caluromys provides an independent test of the arboreal hypothesis (Smith: Annual Report of the Board of Regents of the Smithsonian Institution 1912:553–572, 1913), the visual predation hypothesis (Cartmill: The Functional and Evolutionary Biology of Primates, pp. 97–122, 1972), and the angiosperm exploitation hypothesis (Sussman: American Journal of Primatology, in press) of primate origins. Quantitative data on free-ranging C. derbianus in Costa Rica demonstrate that it is highly arboreal, uses visually directed predation to capture arthropod prey, and makes extensive use of terminal branch foraging, where it feeds on small angiosperm products. These observations are consistent with predictions from each model of primate origins, thus suggesting that the hypotheses are not mutually exclusive but are interdependent. The initial divergence of primates probably involved exploitation of the rich angiosperm products and associated insects found in fine terminal branches; visually directed predation may have evolved as an efficient method of insect capture in the terminal branch milieu.     

Vertical Locomotion in <Emphasis Type="Italic">Micromys minutus</Emphasis> (Rodentia: Muridae): Insights into the Evolution of Eutherian Climbing

Climbing is integral to scansorial and arboreal lifestyles as it enables access to and vertical ranging within the arboreal strata. As early eutherian mammals exhibit osteological correlates for arboreality, it is important to assess the behavioral mechanisms that are related to competent vertical climbing. In this context, we examined climbing gaits in one of the smallest extant rodents, the Eurasian harvest mouse. For these purposes, we filmed six adult Micromys minutus at 240 fps moving on four different substrate sizes (2 mm, 5 mm, 10 mm, 25 mm), during both vertical ascents and descents. All climbing cycles were lateral sequence slow gaits. Upward climbing was characterized by a higher contribution of the hind limbs, longer swing phases, and a significant involvement of stride frequency in velocity modulation. On the other hand, downward climbing was promoted by employing gaits of even lower diagonality, an increased contact with the substrate, enhanced role of the forelimbs, and a subtler modulation of velocity by stride frequency. Eurasian harvest mice effectively negotiate the finest substrates, but their effectiveness decreased significantly on the largest ones. The morphofunctional similarities of M. minutus to Juramaia sinensis and Eomaia scansoria imply analogous behaviors in early eutherians, which apparently contributed to the successful access and exploitation of the fine-branch arboreal milieu. In this way, extant small arboreal mammals can constitute good models for elucidating and comprehending the adaptive significance of behavioral mechanisms that are related to the evolution of arboreality in early mammals.     

A preliminary study of resource overlap between howling monkeys (Alouatta palliata) and other arboreal mammals in the tropical rain forest of Los Tuxtlas,Mexico

Potential resource overlap between howling monkeys and other arboreal mammals was studied in the rain forest of Los Tuxtlas, Veracruz, Mexico. Eight species of mammals belonging to the orders Primates, Carnivora, Rodentia, and Marsupialia were found to share the canopy and to overlap trophically with howling monkeys. These mammals made up 77% and Alouatta 23% of the arboreal mammalian biomass under consideration. The arboreal porcupine and spider monkey were the only mammals that also fed on leaves. However, in this feeding niche, Alouatta is the only important mammalian folivore in Los Tuxtlas, and resource depression derived from leaf-eating insects is more important. The eight arboreal mammals may exert more pressure upon fruit resources, for they consumed 75% of the estimated total dry weight of fruit/ha/yr consumed by arboreal mammals.     

Comparative population structure of three snook species (Teleostei: Centropomidae) from the eastern central Pacific

Three snook species, Centropomus viridis, Centropomus medius, and Centropomus robalito, from the eastern central Pacific, representing three of the four proposed phyletic lineages in the genus, were analyzed for genetic variability by means of allozyme and RAPD to evaluate the divergence between populations at different levels of dispersal ability and to evaluate the importance of barriers to dispersal in the population subdivision and genetic diversity. Levels of genetic diversity among species estimated by allozymes were similar and consistent with the observed levels of differentiation in marine fish species. Mean heterozygosity ranged from 0.089 for C. viridis to 0.10 for C. robalito. Genetic diversity for the snook species studied was slightly higher than the mean estimation reported in allozymes for 106 marine fish (0.055) and for anadromous fish species (0.043 to 0.057). Multilocus allele frequency homogeneity tests and population-subdivision estimates for both allozyme and RAPD markers revealed the existence of population structure in C. viridis and C. medius, in coincidence with geographic separation of samples, whereas no divergence was detected in C. robalito. This finding may be attributed to the greater population size of C. robalito, which originated by a recent population range expansion, and hence the potential for dispersal is mediated by larval drift. Fluctuations in population size and population range expansion are used to explain discrepancies between levels of genetic diversity and population structure in the studied species. Supplementary material to this paper is available in electronic format at     

Ovipositional behaviour and larval performance of Trichoferus campestris (Faldermann) (Coleoptera: Cerambycidae) on cut branches of four North American tree species

1. Velvet longhorned beetle Trichoferus campestris (Faldermann) (Coleoptera: Cerambycidae) is native to Asia. It is reported to be polyphagous in its native range, but hosts have not been screened in its introduced range in the United States.
2. We conducted choice and no-choice host experiments. For choice assays, gravid T. campestris oviposited freely on cut branches of four tree species: Juglans nigra, Malus domestica, Acer saccharum, and Pinus strobus. We investigated oviposition between apple branches with and without cankers. Half of the choice trials underwent winter temperatures.
3. For no-choice assays, eggs were placed on cut branches of these same four species. Half of these branches were later dissected to measure larval development. The rest were exposed to winter temperatures and T. campestris emergence was quantified.
4. Overall, Trichoferus campestris oviposition and development occur more readily on J. nigra and M. domestica than on A. saccharum or P. strobus. The insect oviposited more on cankered than uncankered apple wood. Cold exposure during an overwintering period benefitted insect development in adults.
5. The insect's high degree of polyphagy and ability to reproduce in dry wood has likely facilitated its establishment in multiple locales around the world.

     

Factors Affecting the Compliance and Sway Properties of Tree Branches Used by the Sumatran Orangutan (Pongo abelii)

The tropical arboreal environment is a mechanically complex and varied habitat. Arboreal inhabitants must adapt to changes in the compliance and stability of supports when moving around trees. Because the orangutan is the largest habitual arboreal inhabitant, it is unusually susceptible to branch compliance and stability and therefore represents a unique animal model to help investigate how animals cope with the mechanical heterogeneity of the tropical canopy. The aim of this study was to investigate how changes in compliance and time of oscillation of branches are related to easily observable traits of arboreal supports. This should help predict how supports react mechanically to the weight and mass of a moving orangutan, and suggest how orangutans themselves predict branch properties. We measured the compliance and time of oscillation of branches from 11 tree species frequented by orangutans in the rainforest of Sumatra. Branches were pulled at several points along their length using a force balance at the end of a stiff rope, and the local diameter of the branch and the distance to its base and tip were measured. Compliance was negatively associated with both local diameter and length to the tip of the branch, and positively, if weakly, associated with length from the trunk. However, branch diameter not only predicted compliance best, but would also be easiest for an orangutan to observe. In contrast, oscillation times of branches were largely unaffected by local diameter, and only significantly increased at diameters below 2 cm. The results of this study validate previous field research, which related locomotory modes to local branch diameter, while suggesting how arboreal animals themselves sense their mechanical environment.     

An analysis of chewed food particle size and its relationship to molar structure in the primatesCheirogaleus medius andGalago senegalensis and the insectivoranTupaia glis

The chewed food particle size and shearing capacity of the lower molars of two primate species, the fat-tailed dwarf lemur,Cheirogaleus medius and the bushbabyGalago senegalensis, and an insectivoran, the tree shrew,Tupaia glis, were compared. Differences in the shearing design of the lower molars correlate strongly with the chewed food particle size in these species: the greater the shearing capacity, the smaller the chewed food particles. These three species are of comparable size but differ greatly in diet in the wild.C. medius primarily eats fruit and nectar, whileG. senegalensis andT. glis are largely insect-eaters. The lower molars ofG. senegalensis andT. glis show a much greater shearing capacity than do those ofC. medius. The average length of chewed food particles ofC. medius is significantly larger than that ofG. senegalensis, while that ofT. glis is intermediate between the two primates but is closer to that ofG. senegalensis. Our findings that insect-eating species grind their food more finely than do fruit- and resin-eating species can be correlated with digestibility of foods: finely chewing foods such as fruits which are low in relatively undigestible cell wall components would not greatly improve their digestibility, so a highly efficient food processing apparatus would be less important to the animal's survival. Insect-eaters much more finely chew their foods, implying that there is some constituent of insect bodies difficult to digest, and that grinding increases its digestibility. We suggest that this constituent is chitin.     

Characteristics and use of sleeping sites in Aotus (Cebidae: Primates) in the Amazon lowlands of Peru

In the Amazon lowlands of Peru, Aotus nancymai and A. vociferans were observed to use four different types of sleeping sites: (1) holes in the trunks and branches of dry or senescent trees; (2) concavities in polyaxial branching nodes of trees protected by dense entanglements of creepers, climbing plants, vines, and masses of diverse epiphytes; (3) complex sites among masses of epiphytes, climbers, and vines; and (4) simple sites among thickets and dense foliage. Each type is described. There was competition and sharing of sleeping holes between Aotus and other nocturnal arboreal mammals.     

Developmental Changes in the Sterol Composition and the Glycerol Content of Cuticular and Internal Lipids of Three Species of Flies

The glycerol concentration and the composition of cuticular and internal sterols in three medically and forensically important fly species, viz., Musca domestica, Sarcophaga carnaria, and Calliphora vicina, were analyzed. The cuticular and internal lipid extracts were separated by HPLC‐LLSD, after which the sterol fraction was characterized by GC/MS in total ion current (TIC) mode. The cuticular lipids of M. domestica larvae contained seven sterols, while in pupae and females, six sterols were identified. Five sterols were found in the cuticular lipids of M. domestica males. The internal lipids of M. domestica larvae and pupae contained six and seven sterols, respectively, while those of male and female flies contained only five sterols. Sitosterol, cholesterol, and campesterol were the dominant sterols in M. domestica, while campestanol, stigmasterol, sitostanol, and fucosterol were identified in low concentrations or in traces. In contrast, cuticular and internal lipids of S. carnaria and C. vicina contained only cholesterol. Glycerol was identified in all stages of M. domestica, S. carnaria, and C. vicina. For all the three examined fly species, the present study clearly showed species‐specific developmental changes in the composition of cuticular and internal sterols as well as in the glycerol concentration.     

Is primate-like quadrupedalism necessary for fine-branch locomotion? A test using sugar gliders (Petaurus breviceps)

Locomotor features shared by arboreal marsupials and primates are frequently cited as a functional complex that evolved in the context of a “fine branch niche.” Adaptation to a fine branch niche cannot be understood without considering that small and large arboreal mammals may differ in their biomechanical response to a given branch size. We tested the effects of substrate diameter and orientation on quadrupedal kinematics in a small arboreal marsupial (the sugar glider, Petaurus breviceps). P. breviceps individuals were filmed moving across a flat horizontal surface and on horizontal, inclining and declining poles of diameter 2.5, 1.0, and 0.5 cm. Gait frequencies, limb phases, speeds and duty factors were compared across substrate conditions. P. breviceps had a clear preference for lateral sequence/diagonal couplets gaits, regardless of substrate type, diameter or orientation. Limb phase was significantly influenced by substrate type (higher limb phases on poles vs. the flat surface) and by orientation (higher limb phases on inclined vs. horizontal poles), but was not influenced by pole diameter. Speed was lowest on declines, and duty factors (at a given speed) were highest on the flat board, smallest pole, and on declines. P. breviceps exhibited some parallels, but also some departures from the characteristic patterns of other arboreal marsupials and primates. Notably, limb phase values, on average, remained lower in P. breviceps than those recorded for primates or other arboreal marsupials. We suggest that arboreal mammals of different body sizes may use dissimilar, but apparently equally successful strategies for navigating a “fine branch niche.”     

Distribution of vessel size,vessel density and xylem conducting efficiency within a crown of silver birch (<Emphasis Type="Italic">Betula pendula</Emphasis>)

Spatial patterns in vessel diameter, vessel density and xylem conducting efficiency within a crown were examined in closed-canopy trees of silver birch (Betula pendula). The variation in anatomical and hydraulic characteristics of branches was considered from three perspectives: vertically within a crown (lower, middle and upper crown), radially along main branches (proximal, middle and distal part), and with respect to branch orders (first-, second- and third-order branches). Hydraulically weighted mean diameter of vessels (D _h) and theoretical specific conductivity of the xylem (k _t) exhibited no vertical trend within the tree crown, whereas leaf-specific conductivity of the xylem (LSC_t) decreased acropetally. Variation in LSC_t was governed by sapwood area to leaf area ratio (Huber value) rather than by changes in xylem anatomy. The acropetal increase in soil-to-leaf conductance (G _T) within the birch canopy is attributable to longer path length within the lower-crown branches and higher hydraulic resistance of the shade leaves. D _h, k _t and LSC_t decreased, while vessel density (VD) and relative area of vessel lumina (VA) increased distally along main branches. A strong negative relationship between vessel diameter and VD implies a trade-off between hydraulic efficiency and mechanical stability of xylem. D _h and VD combined explained 85.4% of the total variation of k _t in the regression model applied to the whole data set. Xylem in fast-growing branches (primary branches) had greater area of vessel lumina per unit cross-sectional area of sapwood, resulting in a positive relationship between branch radial growth rate and k _t. D _h, k _t and LSC_t decreased, whereas VD increased with increasing branch order. This pattern promotes the hydraulic dominance of primary branches over the secondary branches and their dominance over tertiary branches. In this way crown architecture contributes to preferential water flow along the main axes, potentially providing better water supply for the branch apical bud and foliage located in the outer, better-insolated part of the crown.     

Angular momentum and arboreal stability in common marmosets (Callithrix jacchus)

Despite the importance that concepts of arboreal stability have in theories of primate locomotor evolution, we currently lack measures of balance performance during primate locomotion. We provide the first quantitative data on locomotor stability in an arboreal primate, the common marmoset (Callithrix jacchus), predicting that primates should maximize arboreal stability by minimizing side-to-side angular momentum about the support (i.e., L_sup). If net L_sup becomes excessive, the animal will be unable to arrest its angular movement and will fall. Using a novel, highly integrative experimental procedure we directly measured whole-body L_sup in two adult marmosets moving along narrow (2.5 cm diameter) and broad (5 cm diameter) poles. Marmosets showed a strong preference for asymmetrical gaits (e.g., gallops and bounds) over symmetrical gaits (e.g., walks and runs), with asymmetrical gaits representing >90% of all strides. Movement on the narrow support was associated with an increase in more “grounded” gaits (i.e., lacking an aerial phase) and a more even distribution of torque production between the fore- and hind limbs. These adjustments in gait dynamics significantly reduced net L_sup on the narrow support relative to the broad support. Despite their lack of a well-developed grasping apparatus, marmosets proved adept at producing muscular “grasping” torques about the support, particularly with the hind limbs. We contend that asymmetrical gaits permit small-bodied arboreal mammals, including primates, to expand “effective grasp” by gripping the substrate between left and right limbs of a girdle. This model of arboreal stability may hold important implications for understanding primate locomotor evolution. Am J Phys Anthropol 156:565–576, 2015. © 2014 Wiley Periodicals, Inc.     

The specialisation of the third metacarpal and hand in arboreal frogs: Adaptation for arboreal habitat?

Skeletal morphology is directly associated with habitat characteristics. To investigate the arboreal adaptation, we studied the osteological morphology of the forelimbs and the third metacarpals of arboreal frogs (Rhacophoridae and Hylidae) and non‐arboreal frogs (Bombinatoridae, Bufonidae, Megophryidae, Ranidae, and Microhylidae) using transparent specimens or X‐rays of skeletons. Our results revealed that the bony knob on the third metacarpal, which formed by a dilated and elongated lateral articular cartilage (AL) through endochondral ossification, occurred only in species of Rhacophorinae. The results of the phylogenetic comparative methods and correlation analysis strongly supported the conclusion that the bony knob is a phylogenetic independent evolution trait and had a significant correlation with the arboreal habitat. Furthermore, anatomical observation showed that a muscle adhered to the bony knob. Therefore, we speculated that the bony knob might act as an enlarged attachment point for larger or more musculatures to help with grasping. In addition, the relative length of the hand showed a significant difference between arboreal and non‐arboreal species (p = .007), suggesting that longer hands might be an arboreal adaptive trait. Overall, this study leads to a deeper understanding of the arboreal adaptation.