Palaeos:		Charadriomorphae
The Vertebrates		Phoenicopteriformes  & Ciconiimorphae

Phoenicopteriformes & Basal Ciconiimorphae

GALLOANSERAE
|
Charadriomorphae 
|--+--Phoenicopteriformes  
|  `--Ciconiimorphae 
|     |--Ciconiiformes
|     |  |--Vulturides 
|     |  `--+--Plataleidae 
|     |     `--Ciconiidae 
|     `--+--Pelecaniformes 
|        `--+--Procellariiformes 
|           `--+--Gaviiformes 
|              `--Sphenisciformes 
`--+--Charadriiformes 
   `--+--Columbiformes 
      `--Psittaciformes 

Taxa on This Page

1. Charadriomorphae
2. Ciconiidae
3. Ciconiiformes
4. Ciconiimorphae
5. Plataleidae
6. Phoenicopteriformes
7. Vulturides

Descriptions

Charadriomorphae: Modern shore birds and an early land bird radiation. Cimolopteryx? (Late Cretaceous fragments), Juncitarsus. 

Range: from the early Eocene, possibly even Late Cretaceous. 

Phylogeny: Galloanserae: Galliformes + Anseriformes + *: (Phoenicopteriformes + Ciconiimorphae) + (Charadriiformes + (Columbiformes + Psittaciformes)). 

Links: Letter on Bird Classification; Charadriomorphae. 011003

Note: It seem increasingly likely that our arrangement is incorrect. ATW090407.

Phoenicopteriformes: flamingos, Juncitarsus, Phoenicopterus (Chilean and American flamingos).  

Range: fr mEc. 

Phylogeny: Charadriomorphae:: Ciconiimorphae + *. 

Characters: Long-legged upside-down filter feeders. Characteristic bent bill; V-shaped upper bill and tongue press into slot in in upper bill to force water out through fine lamellae lining both bills and trap algae and small organisms; larger objects excluded by system of spines and hooks on tongue and upper bill; feathers absent from face; 16-20 cervical vertebrae; middle toenail entire; thin body; flight feathers black; intestinal ceca present; short tail; usually found near alkaline shallow lakes; single-leg stance may be heat-conserving technique; dabbling behavior in larger species as in some ducks and some coraciiforms; often highly colonial; swimming common; ground-nesting in unique mud nests with one egg per clutch; young have two coats of down and are precocial, tended in common creches, but fed by individual parents. 

Links: Phoenicopterus ruber (Greater Flamingo); Eggs - Phoenicopteriformes; flmnh.Flamingo; thewildones.org// flamingo.html; Foto:4902 ; fotos-online.de/ 218.htm. African Savannah: Lesser Flamingo; Birds of Nova Scotia - Greater Flamingo; Virtual Bird Field Guide - Greater Flamingo Phoenicopterus ruber -; Aves: Phoenicopteriformes - Flamingos (German); Phoenicopteriformes; Flameengo (Russian); Bird families; Image List of all Photographed Species w/ Hyperlink; Phoenicopteriformes. 

Image: of Phoeniconias minor from the Oakland Zoo. 020226.

Ciconiimorphae: Most modern shorebirds. 

Range: from the Early Eocene

Phylogeny: Charadriomorphae:: Phoenicopteriformes + *: Ciconiiformes + (Pelecaniformes + (Procellariformes + (Gaviiformes + Sphenisciformes))). 

Links: Neornithes -- The Dinosauricon; "Ciconiimorphae" after Olson, 1985, Ericson, 1995 and Feduccia, ...; 

Note: This group is a an island of comparative stability in the tectonically hyper-active area of bird relationships. This may only make it a more attractive target in future, since there is nothing yet that really compels us to accept this taxon as monophyletic. ATW030930.

Ciconiiformes: storks, herons, egrets, ibis, etc. 

Range: fr lwEc. Ciconiimorphae: (Pelecaniformes + (Procellariformes + (Gaviiformes + Sphenisciformes))) + *: Vulturides + (Plataleidae + Ciconiidae). 

Characters: Stork-like forms, mostly large wading birds with large bills; only a single pair of sternotracheal muscles in the syrinx; 16-20 cervical vertebrae; diastataxic (fifth secondary feather absent, but 5th secondary covert is present); feet not webbed; middle ungual laterally expanded, (pectinate in some families); intestinal ceca present, nearly always minute; largely fresh water or terrestrial habitat; not filter feeders, mostly feed on fish, crustacea, insects, carrion; do not swim for food; northerly species migrate; most nest in trees; strong flyers with broad wings; altricial young; generally colonial, but use of sound unusual; social communication by displays and "rituals." 

Links: Ciconiiformes Order; World Birds Taxonomic List: Genera and species with citations.; Ciconiiformes (really gorgeous photographs); Ciconiiformes (in Spanish); Otryad Anstoobrazhnie (Russian). 

Discussion: In most cultures, at least some members of the Ciconiiformes -- herons, storks, ibis, egrets and so on -- have always had an unusual status as objects of religious or artistic veneration. The attraction is certainly not phylogenetic. The morphologically similar, but unrelated, cranes are treated in the same way. Sometimes hauntingly beautiful evocations of these birds are known from antiquity in, for example, Greece, Africa, Egypt, China and Japan. Certainly the vulturid condors have influenced the art and religion of the American Pacific coast from Alaska to Chile. In Indo-European cultures, by contrast, the long-legged water bird tends to be a comical or even evil figure. The cultural phylogeny of these symbols would be an interesting study in itself, and their relative stability over millennia is surprising.

Certainly one of the reasons for the attention given the ciconiiforms is their strange and alien way of moving. That special style of movement, whether perceived as graceful or comically awkward, is emphasized in the behavioral rituals common in the group. Some ciconiiforms are completely silent, and vocalization in most species is fairly limited. Thus, rituals and displays are the primary means of communication.

The study of avian behavior has itself been an interesting example of human behavior and the not-quite clean line between science and political fashion. Much of Konrad Lorenz's original work on imprinting in the 1940's and 1950's concerned bird behavior. This coincided with Skinner's development of the behavioral approach to psychology. For various reasons, this body of research was picked up by a number of different political factions as either tending to prove or disprove their world views, politics and economic theories, and the scientific and political threads in this area have become hopelessly entangled. It is easy to deprecate this sort of thing, but hard to avoid it. The very success of science has taught us to respect scientific truth. We want our beliefs to be scientifically verifiable. That cannot be a bad thing. Yet, beliefs are tools for simplifying and dealing with an infinitely complex universe. They are, by their nature, not entirely rational. We inevitably tend to interpret our science based on our beliefs about the universe in which the experiment takes place -- but the two will never quite match.

As a result of some of the friction at this interface between politics and science, serious work in this area has been a little light over the last couple of decades. After all, what serious ornithologist wants to be condemned as a racist or a communist because of some innocent comment about the mating behavior of whooping cranes? As silly as it may seem, this sort of thing happens when one touches near deeply-held religious or philosophical beliefs, and it cannot help an aspiring researcher in the highly competitive world of grants and tenure-track appointments. Against this background, it is a rare treat to see a paper such as Beth Slikas' treatment of ciconiid behavior and phylogeny. Slikas (1998). It is even more heartening to see a relatively junior post-doctoral researcher, especially one who writes so well, bring some fresh ideas to this difficult area.

There is nothing radical about Slikas's paper -- and hopefully nothing of political significance to anyone. Her thesis work at the University of Pennsylvania seems to have been related DNA molecular phylogenetics. She is currently at the National Zoo and the Smithsonian working on ancient DNA. In the course of her thesis work, she apparently came across the extensive body of comparative behavioral observations on the courtship displays of storks compiled by M. Phillip Kahl in the 1960's and 70's. Fortunately, these displays are rather stereotyped and show little intraspecific variation. Thus, it was possible for Slikas to code the displays as if they were molecular or morphological characters and analyze them on PAUP.

For those with a specialized interest, Slikas found 24 equally parsimonious trees. Thus, the results are somewhat ambiguous, but generally look (to the genus level) like this:

CICONIIDAE
|--Leptoptilos
|--+--+--Ephippiorhynchus
|  |  `--Jabiru
|  `--Ciconia
`--+--Mycteria
   `--Anastomus

This was entirely consistent with Slikas's molecular work, except that the latter had unambiguously placed Leptoptilos as the sister of the Mycteria + Anastomus clade. The more generally applicable result is Slikas's suggestion that there seems to be a hierarchy of homoplasy. Specifically, rituals associated with initial mate selection, such as the male's "advertising" of his nest site and the female's expression of interest in the male's real estate, were quite stable. Behaviors related to later events, such as copulation and pair-bond affirmation were more phylogenetically plastic. Finally, behaviors not related to mating, such as the "anxiety stretch" or aggression displays, were quite variable.

One must be very careful not to over-interpret results in this area; but this is clearly a very interesting observation and deserves some follow-up. Other work, reviewed in Düttmann et al. (1998), suggests that initial sexual displays can be distorted in grossly abnormal developmental situations, such as when an individual is raised by parents of a different species. However, even hatchlings raised in isolation eventually arrive at the "correct" behaviors, so the pathological case may mean rather little. What is more curious is that avian songs seem to be quite plastic and are heavily influenced by imitation and learning. Songs are clearly the best-known and most obvious sexual display and mate-selection behaviors of passerine birds. Thus one can tentatively conclude that variability in this device has some selective advantage, perhaps by communicating metabolic health or encouraging a healthy level of exogamy. Yet the corresponding behaviors of storks seem to have been relatively stable for periods on the order of tens of millions of years.

There are any number of possible explanations for these contrasting observations. Passerines are not colonial in the same way as some storks. There may be stable and variable elements of birdsong which need to be separated out. The costs of mating "error" may be higher in storks for reasons relating to size or developmental limitations. The difference may simply be that passerines are significantly brighter than storks and less likely to be confused by variation. Whatever the case, the issues raised by this work should attract some further -- but, one hopes, discreet -- scientific attention. ATW 010114

References: Düttman et al. (1998), Slikas (1998). 010110.

Vulturides (= Cathartidae): teratorns & New World vultures, condors. 

Phylogeny: Ciconiiformes: (Ciconiidae + Plataleidae) + *. 

Characters: Large soaring birds. Bill hook-like; nostrils externally perforate; holorhinal; palate indirectly desmognathous; maxillopalatines widely separated but co-ossify with medial nasal septum; maxillopalatines in form of scroll-like plates; vomer absent; pterygoids twisted; basipterygoid process present; unfeathered usually brightly colored heads (adults); sternum entire (not notched on posterior surface); legs stout; toes thin; claws short; hallux small and elevated; basal web between toes 2 and 3; black or dark brown plumage; scavengers; generally voiceless. 

Links: GeoZoo: Herons, Storks & Relatives! (Ciconiiformes); Bird families; Condor Skull Replicas Offered by Skulls Unlimited; Birds - Everything About Vultures and Condors; Vulturides; FLMNH bird photos archive page 12; Field Studies in Biology: Bird Identification; Order Falconiformes; vulture; Vulture's Purpose; New World Vultures; Cathartidae (Portugese); Family Cathartidae; Première partie: Etude bibliographique (French -- neornithine interordinal relationships). 

Image: Turkey vulture, Cathartes aura, from Dr. Dan Sudia, by permission. 010313.

Plataleidae (= Threskiornithidae?): Ibis & spoonbills (Threskiornis, Platalea, etc.). 

Range: Fr mEc. 

Phylogeny: Ciconiiformes:: Ciconiidae + *. 

Characters: Medium to large wading birds with long legs, with a long, slender, grooved, and decurved bill, or (spoonbills) with a flattened spatulate bill; crests common; long neck; middle ungual entire, digits may be slightly webbed; halux raised; not pectinate; plumage variable--white, brown, black (pink or red in two tropical species). 

Links: Ciconiiformes; Threskiornithidae Family; GeoZoo: Herons, Storks & Relatives! (Ciconiiformes); Plataleidae [Threskiornithidae]; Foto-album: Plataleidae; Foto-album- Plataleidae.  ATW030117.

Ciconiidae: storks, e.g. Balaeniceps, Leptoptilos, Mycteria. 

Range: from the Late Eocene (Early Oligocene?) worldwide. 

Phylogeny: Ciconiiformes:: Plataleidae + *. 

Characters: Long - legged, stocky birds, 60-150 cm; head and neck sparsely feathered or bare; long, stout bill, sometimes curved distally and usually ungrooved; long, broad wings; tail short; partial web between middle and outer toes, halux smaller and raised; remiges usually black & remainder black or white; no sexual dimorphism; fly with head & legs extended; many feed on land (carrion) or aquatic animals in shallow or temporary waters; poorly developed vocalizations (some use bill clattering); most colonial; biparental care; biparental construction of large, platform nest. 

Image: Wood stork, Mycteria americana, from Dr. Dan Sudia, by permission. 

Links: GeoZoo: Herons, Storks & Relatives! (Ciconiiformes); Ciconiidae Family; Aves: Ciconiidae- Stoerche; Semeistvo Aestovye (Russian); Dierentuin.Net Dieren Database Ooievaars (Dutch); Stork Skull Replicas Offered by Skulls Unlimited; A Sibley-Monroe-like list of World Birds;  ciconiiformes.html; Florida Nature- Ciconiidae -; Ciconiidae; Aves- Ciconiidae- Stoerche; Ciconiidae; APUS.RU | Семейство ·Аистовые - Ciconiidae.    

References: Slikas (1998). ATW030611.