Prey capture and accommodation in the sandlance, Limnichthyes fasciatus (Creediidae; Teleostei)

The eyes of the sandlance, Limnichthyes fasciatus (Creediidae, Teleostei) move independently and possess a refractive cornea, a convexiclivate fovea and a non-spherical lens giving rise to a wide separation of the nodal point from the axis of rotation of the eye much like that of a chameleon. To investigate this apparent convergence of the visual optics in these phylogenetically disparate species, we examine feeding behaviour and accommodation in the sandlance with special reference to the possibility that sandlances use accommodation as a depth cue to judge strike length. Frame-by-frame analysis of over 2000 strikes show a 100% success rate. Explosive strikes are completed in 50 ms over prey distances of four body lengths. Close-up video confirms that successful strikes can be initiated monocularly (both normally and after monocular occlusion) showing that binocular cues are not necessary to judge the length of a strike. Additional means of judging prey distance may also be derived from parallax information generated by rotation of the eye as suggested for chameleons. Using photorefraction on anaesthetised sandlances, accommodative changes were induced with acetylcholine and found to range between 120 D and 180 D at a speed of 600–720 D s⁻¹. The large range of accommodation (25% of the total power) is also thought to be mediated by corneal accommodation where the contraction of a unique cornealis muscle acts to change the corneal curvatures. Accepted: 8 December 1999     

Terrestrial optics in an aquatic eye: The sandlance,Limnichthytes fasciatus (Creediidae,Teleostei)

The sandlance, Limnichthyes fasciatus (Creediidae, Teleostei), behaves like a marine chameleon, with independent movements of its turret-like eyes, highly-effective camouflage and rapid strikes for isolated, mobile prey at close quarters. The optical system has a fixed circular pupil, a deep pit fovea and a flattened lens unlike any other teleost lens so far described. The convex, laminated structure of the cornea is also unparalleled in a teleost which suggests that the cornea may play a refractive role that might compensate for the reduced power of the flattened lens. This suggestion has been supported in the present investigation by four independent sets of observations:- i. Purkinje images formed underwater by the cornea; ii. Measurements of the magnification of intra-corneal iridophores viewed through the corneal lenticle; iii. Measurements of the magnification produced by the dissected corneal lenticle and lens when viewed over a grating; iv. Ray tracing experiments comparing the degree of refraction produced by the lens and by the corneal lenticle. All experimental observations confirm that the cornea of the sandlance has a significant refractive role, with a power of approximately 200 D compared with a power of 550 D for the lens. This is the first report of a significant refractive role played by the cornea in a teleost. The optical system of lens plus cornea, in combination with a deep pit fovea, may be more suitable for the detection and visual localisation of small, moving, underwater prey than the conventional wide-field spherical lens system of other teleosts. The evolutionary convergence of this marine optical system and lifestyle with those of the chameleon is remarkable, given the constraints imposed by underwater optics.     

Binocular vision and accommodation in prey-catching chameleons

We have studied the role of accommodation and binocular convergence in the predatory behaviour of two chameleon species (Chamaeleocalyptratus, C. dilepis). Accommodation measurements support earlier observations that accommodation is the major distance cue. Specifically, accommodation speed (60 D s⁻¹), amplitude (45 D) and precision (no significant under-accommodation) were superior to those of other terrestrial vertebrates. Similar to other vertebrates, accommodation was accompanied by a prominent pupillary constriction (pupillary near response). Accommodation could be coupled or uncoupled in both eyes, depending on the experimental situation or the phase of the predatory behavioural sequence. Uncoupled accommodation occurred: 1. During scanning saccadic eye movements for prey detection. Only one eye accommodated appropriately, the other adopted a hyperopic resting refractive state. Attention switched from one eye to the other at approximately 1-s intervals. 2. During initial stages of distance estimation. Coupled accommodation only occurred shortly before the tongue shot. Coupling was demonstrated by either covering one eye with a lens or covering one eye with an infrared light transmitting cut-off filter which still permitted refraction to be measured. In both cases the amount of accommodation was identical in both eyes. Search-coil measurements showed that the angle of convergence of both eyes is too variable to permit triangulation or to provide the basic requirement for stereopsis (matching corresponding points). We conclude that coupling of accommodation serves to improve accommodation precision rather than to permit stereopsis. Accepted: 3 September 1997     

Visual fields of orb web and single line web spiders of the family Uloboridae (Arachnida,Araneida)

Summary In the family Uloboridae, web reduction is associated with changes in web monitoring posture and prosomal features. A spider must extend its first pair of legs directly forward to monitor the signal line of a reduced web. This posture is facilitated by shifts in prosomal musculature that cause reduced web uloborids to have a narrower anterior prosoma, a reduced or absent anterior eye row, and prominent posterior lateral eye tubercles. The eye tubercles and larger posterior eyes of these uloborids suggest that web reduction may also be accompanied by ocular changes that compensate for reduction of the anterior eyes by expanding the visual fields of the posterior eyes. A comparison of the visual fields of the eight-eyed, orb web species Octonoba octonaria and a four-eyed, reduced web Miagrammopes species was made to determine if this is true. Physical and optical measurements determined the visual angles of each species' eyes and the pattern of each species' visual surveillance. Despite loss of the anterior four eyes, the Miagrammopes species has a visual coverage similar to that of O. octonaria. This is due to (1) an increase in the visual field of each of the four remaining Miagrammopes eyes, accruing from an extension of the retina and an increase in the lens' rear radius of curvature, and (2) a ventral shift of each visual axis, associated with the development of an eye tubercle and an asymmetrical expansion of the retina. Miagrammopes monitor their simple webs from twigs or moss where they are vulnerable to predation. Therefore, maintenance of visual cover may enable them to detect predators in time to assume or maintain their characteristic, cryptic posture. It may also allow them to observe approaching prey and permit them to adjust web tension or prepare to jerk their webs when prey strikes.     

Morphological characteristics of eyes and retinas of two sardines (Sardinops melanostictus and Etrumeus sadina,Clupeidae) and an anchovy (Engraulis japonicus,Engraulididae)

The morphology of the eyes and distribution of retinal ganglion cells in two sardine species (Sardinops melanostictus and Etrumeus sadina, Clupeidae) and the Japanese anchovy (Engraulis japonicus, Engraulididae) were investigated anatomically and histologically. The eyes of the sardines faced a slightly dorsolateral direction with the visual field extended obliquely upward. In contrast, the eyes in the anchovy were almost laterally directed. It was hypothesized that the sardines may have an advantage in receiving more downward irradiance compared with the anchovy. The lens muscle was larger in these three species than in many other teleosts, and its surface was entirely melanin‐pigmented. Also, the lens muscle directly and tightly adhered to the backside surface of the iris. The relative area of the lens muscle to the area of the lens, a referential value of the relative power of visual accommodation were notably larger in the species studied than in other teleost values that have been previously reported. A higher M/L% value of these clupeid fishes could facilitate fast and wide ranging visual accommodation and was considered to be associated with maintaining and/or re‐establishing school formations quickly. Analysis of topographical distributions of cells in the ganglion cell layer showed that cell density was highest in the ventrotemporal quadrant of the retina (temporal of the optic cleft) in all three species. Another potentially important role for the black‐pigmented lens muscle may be to block the specialized retinal area from intense sunlight that scatters and irradiates upward or laterally in the surface waters that they inhabit. Thus, the sardine and anchovy may take advantage of efficient detection of visual signals in the frontal‐upward direction and further improve visibility of the target in this direction. J. Morphol. 276:415–424, 2015. © 2014 Wiley Periodicals, Inc.     

Optical properties of a cephalopod eye (the short finned squid,Illex illecebrosus)

Summary The extent to which the cephalopod eye is optically similar to the teleost eye was determined by measuring refractive error, accommodative ability, spherical and chromatic aberrations, and refractive indices and radii of curvature of the ocular media. The squid eye is well corrected optically underwater although a tendency toward hyperopia exists. This may be due to the existence of chromatic aberration and the fact that an aquatic environment is somewhat limited to the blue end of the spectrum. Accommodation takes place by movement of the lens toward the retina in a manner similar to the teleost eye. However, the squid lens is not spherical but slightly flattened. The lens is overcorrected as far as spherical aberration is concerned. Thus peripheral light rays focus further from the lens than paraxial ones. The function of this unusual example of lens development is unknown.     

Accommodation behaviour during prey capture in the vietnamese leaf turtle (<Emphasis Type="Italic">Geoemyda spengleri</Emphasis>)

Vietnamese leaf turtles (Geoemyda spengleri) were tested for their ability to focus on prey objects at various distances. Accommodation was continuously measured by infrared photoretinoscopy. All animals investigated during this study showed a surprisingly high precision of accommodation over a range of over 30 D. Measured accommodation matched the target distance accurately for distances between 3 and 17 cm. The turtles switched between independent and coupled accommodation in the two eyes. Independent accommodation was observed when the turtles inspected their environment visually without a defined object of interest. Coupled accommodation was only observed during binocular prey fixation. When a turtle aimed at a target, the symmetrical focus of both eyes persisted even if vision was totally blocked in one eye or altered by ophthalmic lenses. This suggests that the eyes were linked by internal neuronal mechanisms. The pupil of the eye responded clearly to changes in ambient light intensity. A strong decrease in pupil size was also observed when the eye was focused on a close target. In this case, the constriction of the pupil probably aids in the deformation of the eye lens during near-accommodation.     

Ecomorphology of the eyes and skull in zooplanktivorous labrid fishes

Zooplanktivory is one of the most distinct trophic niches in coral reef fishes, and a number of skull traits are widely recognized as being adaptations for feeding in midwater on small planktonic prey. Previous studies have concluded that zooplanktivores have larger eyes for sharper visual acuity, reduced mouth structures to match small prey sizes, and longer gill rakers to help retain captured prey. We tested these three traditional hypotheses plus two novel adaptive hypotheses in labrids, a clade of very diverse coral reef fishes that show multiple independent evolutionary origins of zooplanktivory. Using phylogenetic comparative methods with a data set from 21 species, we failed to find larger eyes in three independent transitions to zooplanktivory. Instead, an impression of large eyes may be caused by a size reduction of the anterior facial region. However, two zooplanktivores (Clepticus parrae and Halichoeres pictus) possess several features interpreted as adaptations to zooplankton feeding, namely large lens diameters relative to eye axial length, round pupil shape, and long gill rakers. The third zooplanktivore in our analysis, Cirrhilabrus solorensis, lacks all above features. It remains unclear whether Cirrhilabrus shows optical specializations for capturing planktonic prey. Our results support the prediction that increased visual acuity is adaptive for zooplanktivory, but in labrids increases in eye size are apparently not part of the evolutionary response.     

Modulation of prey capture kinematics in the cheeklined wrasse Oxycheilinus digrammus (Teleostei: Labridae)

The ability to modulate prey capture behaviors is of interest to organismal biologists as it suggests that predators can perceive features of the prey and select suitable behaviors from an available repertoire to successfully capture the item. Thus, behavior may be as important a trait as morphology in determining an organism's diet. Using high-speed video, we measured prey capture kinematics in three cheeklined wrasse, Oxycheilinus digrammus. We studied the effects of three experimental prey treatments: live fish, dead prawn suspended in the water column, and dead prawn pieces anchored to the substrate in a clip. Live prey elicited significantly more rapid strikes than dead prey suspended in the water column, and the head of the predator was expanded to significantly larger maxima. These changes in prey capture kinematics suggest the generation of more inertial suction. With greater expansion of the head, more water can be accelerated into the buccal cavity. The attached prey treatment elicited strikes as rapid as those on live prey. We suggest that the kinematics of rapid strikes on attached prey are indicative of attempts to use suction to detach the prey item. More rapid expansion of the buccal or mouth cavity should lead to higher velocities of water entering the mouth and therefore to enhanced suction. Further modulation in response to the attached prey item, such as clipping or wrenching behaviors, was not observed. J. Exp. Zool. 290:88-100, 2001.     

Visual optics: The sandlance eye breaks all the rules

The eyes of the sandlance differ from those of other fish, both optically and in the kinds of movements they make. The predatory behaviour of these tiny fish not only makes their lifestyle similar to that of a chameleon, but has led to several extraordinary examples of convergence in the visual system.     

Development of the compound eyes of dragonflies (Odonata). III. Adult compound eyes

The distribution of ommatidial diameters and interommatidial angles, as determined by measuring the angles between the optic axes of adjacent ommatidia, are mapped across the surface of the compound eyes of a variety of species selected for different adult behaviors, developmental histories, and taxonomic positions. The size of the visual fields, prey capture foveas, foveas composed of large dorsal ommatidia, and other specializations in the numbers of ommatidia that view various directions in the visual field are discussed in relation to adult behavior. Advanced species have less resemblance between their larval and adult eyes than primitive species. In contrast to their larvae, adults increase the monocular resolution of each eye at the expense of binocular vision. Most species have foveas which view in approximately the anterior direction, instead of in a region of binocular overlap, and many species have foveal bands which view along the horizon. Some advanced perching species, which approach their prey and other odonates from below, have an additional vertical foveal band that views along a vertical plane from the anterior direction to a more dorsal direction. The most unusual foveal band is seen in active flying species. The large dorsal ommatidia of the migratory Anax junius, which cover approximately one third of the eye surface, view a narrow region of the visual field that extends along a plane from the most lateral direction of one eye to a dorsal direction, and continues without interruption to the most lateral direction of the other eye.     

Visual accommodation and active pursuit of prey underwater in a plunge-diving bird: the Australasian gannet

Australasian gannets (Morus serrator), like many other seabird species, locate pelagic prey from the air and perform rapid plunge dives for their capture. Prey are captured underwater either in the momentum (M) phase of the dive while descending through the water column, or the wing flapping (WF) phase while moving, using the wings for propulsion. Detection of prey from the air is clearly visually guided, but it remains unknown whether plunge diving birds also use vision in the underwater phase of the dive. Here we address the question of whether gannets are capable of visually accommodating in the transition from aerial to aquatic vision, and analyse underwater video footage for evidence that gannets use vision in the aquatic phases of hunting. Photokeratometry and infrared video photorefraction revealed that, immediately upon submergence of the head, gannet eyes accommodate and overcome the loss of greater than 45 D (dioptres) of corneal refractive power which occurs in the transition between air and water. Analyses of underwater video showed the highest prey capture rates during WF phase when gannets actively pursue individual fish, a behaviour that very likely involves visual guidance, following the transition after the plunge dive's M phase. This is to our knowledge the first demonstration of the capacity for visual accommodation underwater in a plunge diving bird while capturing submerged prey detected from the air.     

Predator-specific camouflage in chameleons

A crucial problem for most animals is how to deal with multiple types of predator, which differ in their sensory capabilities and methods of prey detection. For animals capable of rapid colour change, one potential strategy is to change their appearance in relation to the threat posed by different predators. Here, we show that the dwarf chameleon, Bradypodion taeniabronchum, exhibits different colour responses to two predators that differ in their visual capabilities. Using a model of animal colour perception to gain a 'predator's eye view', we show that chameleons showed better background colour matching in response to birds than snakes, yet they appear significantly more camouflaged to the snake visual system because snakes have poorer colour discrimination.     

Visual optics in toads (Bufo americanus)

Aspects of visual optics were investigated in the American toad (Bufo americanus). The development of the refractive state of the eye during metamorphosis was followed with IR photoretinoscopy. Frozen sections documented the changes in optical parameters before and after metamorphosis. There is a difference in light sensitivity between juvenile and adult toads. Binocular accommodation in adult toads was observed. 1. IR photoretinoscopic measurements showed that the refractive state of the eye changed very rapidly during metamorphosis, about 10 D/h while the animal entered the terrestrial habitat. 2. Frozen sections showed that the almost spherical lens in a tadpole eye had flattened in a just metamorphosed toad's eye while at the same time the distance of the lens to the retina had decreased. However, the morphological measurements were not sufficiently sensitive to record the relatively small changes in ocular dimensions that were responsible for the rapid changes in refractive state during metamorphosis. 3. Schematic eyes, with homogeneous and non homogeneous lenses, were constructed for tadpoles, juvenile toads, and adult toads. 4. Nonparaxial raytracing studies in schematic eyes suggested that the lenses of animals of the three developmental stages tadpole, juvenile toad, and adult are not homogeneous but have a refractive index gradient. The raytracing studies indicated that the refractive index gradient is different for the different developmental stages, being highest in the tadpole lens. 5. The observations of toads during feeding behavior at different light levels showed an increased light sensitivity in the adult nocturnal toads in contrast to the juvenile animals, which are diurnal. The increased light sensitivity could partly be explained with an increase in aperture and an increase in red rod outer segments. To fully explain the higher light sensitivity in adult toads, changes in neuronal parameters had to be assumed. 6. Retinoscopic measurements of the resting refractive state in the adult toad showed a hyperopic defocus of about +8 D. By subtracting the measurement artefact for retinoscopy, the true resting focus was found to be nearly emmetropic. 7. The amount of natural accommodation in adult toads during normal feeding behavior was investigated with IR photoretinoscopy. Binocular accommodation of about 8 D was observed.     

QUANTIFYING ERRORS ASSOCIATED WITH USING PREY SKELETAL STRUCTURES FROM FECAL SAMPLES TO DETERMINE THE DIET OF STELLER'S SEA LION (EUMETOPIAS JUBATUS)

We examined the digestion and passage times of bones and other hard parts from pollock, herring, salmon, and sandlance recovered from two juvenile captive Steller's sea lions ( Eumetopias jubatus ) subjected to varying activity levels. Key bones that could be identified to species were distributed over an average of 3.2 scats (range 1–6) following a single meal, with pollock remains occurring in significantly more scats than other species. Relying on otoliths alone to determine the presence of prey resulted in significantly fewer prey being identified than if other structures were also used (such as vertebrae, jaw bones, and teeth), particularly for salmon. Using either technique, there were significant differences in the likelihood that bones would be recovered from the series of scats produced following a meal, with pollock recovery exceeding herring (by three-fold) and sandlance (by eight-fold). Differences between species were reduced when recovery was calculated on a per scat basis rather than over multiple scats. Active animals passed greater numbers of bones, but the overall effect on prey recovery estimates was not significant. Defecation times of prey structures from a meal were variable and ranged from an initial 2–56 h to a final 28–148 h. The time interval to pass 95% of recovered structutes varied by a factor of two among prey species, and was highest for pollock due to retention beyond 65 h.     

Allometry and resolution of bee eyes (Apoidea)

A sample of compound eyes from 15 species of female pollen foraging bees (apiform Apoidea) was morphometrically analyzed. These species were chosen for size differences, different social organization, and a wide geographic and taxonomic distribution (Apidae, Megachilidae, Andrenidae, Halictidae). The results demonstrate the following characteristics for the typical compound eye in female foraging bees: (1) the vertical diameter of the eye is about twice the horizontal diameter; (2) the eyes of diurnal foragers scale isometrically with body size; (3) the eyes of three species of nocturnal foragers have about 1.8 times the surface area as compared to diurnal foragers of matching size; (4) the number of ommatidia per eye range from about 1000 in Perdita minima to about 16 000 in Xylocopa latipes; and (5) the corresponding mean interommatidial angles range from 4.7 to 1.2 degrees . Body size, rather than species-specific ecological adaptation, is the major (97%) determinant of the number of ommatidia per eye in diurnal, as well as nocturnal foragers. The number of ommatidia per eye, and hence the visual resolution, is proportional to the square root of both body size and eye size across all species studied. The eye parameter (the product of the mean interommatidial angle and the mean lens diameter) increases slightly with decreasing body size. All this is taken as evidence that the features of the bees' visual macro-niche remained largely constant over the roughly 130 million years of their macro-evolution.     

The comparative morphology and evolution of the eyes of caecilians (Amphibia,Gymnophiona)

Summary Caecilians (Amphibia, Gymnophiona) have been reported to have vestigial eyes, to lack some or all of the extrinsic eye muscles and their nerves, and to utilize eye muscles and glands, or derivatives of them, to effect movement of the tentacle, a chemosensory structure unique among vertebrates. Morphological evidence indicates that the eye is a functional photoreceptor in virtually all species examined, with an intact retina and optic nerve. The pattern of retention of extrinsic muscles varies. The ontogeny of the eye of Dermophis mexicanus is typical of that of most vertebrates, though components of accommodation never develop. Several taxa are reported in the literature to lack various eye structures; the present study reveals them to be variously present. Evolutionary trends in caecilian eye morphology include the following: (1) the eye is overlain by thicker, often glandular skin, to overlain by bone as well as skin; (2) extrinsic muscles become attenuate, and some to all may be lost; (3) the retina has the typical vertebrate layered organization, to having a reduced cell number, to becoming net-like rather than stratal; (4) the optic nerve is present, becoming attenuate, perhaps represented only by glial cells; (5) the lens is round (aquatic forms, larval and adult) to spheroid; lens crystalline to cellular (retention of the embryonic condition) to amorphous to absent; (6) the vitreous body is reduced or lost; (7) the cornea adheres to the overlying dermis or periosteum; the lens is free to adherent to cornea to adherent to both cornea and retina. Scolecomorphids have the eye pulled out of the socket and embedded in the tentacle under the skin of the upper jaw. This pattern of trends in eye reduction is similar to that observed in other vertebrate lineages that are fossorial or troglobitic.     

Specialized ommatidia for polarization vision in the compound eye of cockchafers,Melolontha melolontha (Coleoptera,Scarabaeidae)

Summary The superposition eye of the cockchafer, Melolontha melolontha, exhibits the typical features of many nocturnal and crepuscular scarabaeid beetles: the dioptric apparatus of each ommatidium consists of a thick corneal lens with a strong inner convexity attached to a crystalline cone, that is surrounded by two primary and 9–11 secondary pigment cells. The clear zone contains the unpigmented extensions of the secondary pigment cells, which surround the cell bodies of seven retinula (receptor) cells per ommatidium and a retinular tract formed by them. The seven-lobed fused rhabdoms are composed by the rhabdomeres of the receptor cells 1–7. The rhabdoms are optically separated from each other by a tracheal sheath around the retinulae. The orientation of the microvilli diverges in a fan-like fashion within each rhabdomere. The proximally situated retinula cell 8 does not form a rhabdomere. This standard form of ommatidium stands in contrast to another type of ommatidium found in the dorsal rim area of the eye. The dorsal rim ommatidia are characterized by the following anatomical specializations: (1) The corneal lenses are not clear but contain light-scattering, bubble-like inclusions. (2) The rhabdom length is increased approximately by a factor of two. (3) The rhabdoms have unlobed shapes. (4) Within each rhabdomere the microvilli are parallel to each other. The microvilli of receptor 1 are oriented 90° to those of receptors 2–7. (5) The tracheal sheaths around the retinulae are missing. These findings indicate that the photoreceptors of the dorsal rim area are strongly polarization sensitive and have large visual fields. In the dorsal rim ommatidia of other insects, functionally similar anatomical specializations have been found. In these species, the dorsal rim area of the eye was demonstrated to be the eye region that is responsible for the detection of polarized light. We suggest that the dorsal rim area of the cockchafer eye subserves the same function and that the beetles use the polarization pattern of the sky for orientation during their migrations.     

Visual-optical characteristics of caged and semifree-ranging monkeys

Five species of macaques and one species of mangabey comprising a total of 324 monkeys (132 males and 192 females) were given visual examinations which included measures of corneal curvature, depth of the anterior chamber, thickness of the lens, depth of the vitreous chamber, total axial length, total power of the eye and intraocular pressure as well as the refractive error of eye under cycloplegia in the anesthetized monkey. Cercocebus and Macaca mulatta demonstrated the greatest amount of myopia and Macaca fascicularis the least, with M. speciosa, M. fuscata, and M. nemestrina in order between these extremes. The M. nemestrina eye approximates that of the chimpanzee in size and relationship of the components. The findings indicate that males have larger eyes than females, the curvature of the cornea decreases as the axial length increases, and there is a close relationship between the length of the eye and the amount of myopic refractive error. There is some evidence that amount of myopia and the percentage of animals showing a myopic refractive error are related to the visual conditions and the behavior patterns of the different species.     

Comparative analysis of the size and shape of the lizard eye

Lizards occupy both scotopic (light-limited) and photopic (light-rich) environments, thereby making this clade ideal for analyses of eye morphology adaptations. This study examines how in lizards the morphology of the eye varies according to activity in these different light environments. Measurements were collected on corneal diameters and axial lengths of the eye for 239 specimens of 116 lizard species (including Sphenodon) that include both species with scotopic and photopic visual adaptations. I show that the light level available to a lizard for vision has a significant effect on eye shape and size. Scotopic lizards have eye shapes that are optimized for visual sensitivity, with larger corneal diameters relative to axial lengths. However, photopic lizards do not exhibit absolutely larger axial lengths than do scotopic lizards, and the groups have the same absolute axial lengths of the eye. Results also indicate that the light level the lizard functions under is a more significant influence on eye shape, as defined by the relationship between corneal diameter and axial length of the eye, than is phylogeny.