Zander, R. H. 2019. Macroevolutionary Evaluation

Zander, R. H. 2019. Macroevolutionary evaluation methods extended, consolidated, and exemplified with Anoectangium (Pottiaceae, Bryophyta) in North America and the Himalayas. Annals of the Missouri Botanical Garden 104: 324-338.

The original paper is available through the Annals of the Missouri Botanical Garden. This version for distribution has the same content and pagination.

ABSTRACT Methods of macroevolutionary systematics as recently modified are explained and applied to the genus Anoectangium Schw¨agr. (Pottiaceae) in hyperoceanic areas of coastal northwestern North America. It was revealed that this area harbored species like those reported for the Himalayan region of northern India. Keys are provided for North American and Himalayan species of the genus. A macroevolutionary analysis, detailed in the Methods section, distinguishes and gives relationships between the species. There is strong Bayesian support for progenitor-descendant pairs and lineages. Anoectangium thomsonii Mitt. is considered a synonym of A. aestivum (Hedw.) Mitt. Anoectangium crassinervium Mitt. is transferred to Molendoa Lindb., under a nomen novum with A. handelii Schiffn. as a synonym. Anoectangium incrassatum Broth., related to the Asian A. clarum Mitt., is reported as a well-characterized species from the West Indies. Anoectangium stracheyanum Mitt. is re-lectotypified. Distinctions between A. aestivum and A. euchloron (Schw¨agr.) Spruce in the New World are clarified. Causal explanations in systematics are equated with entropy maximization in Shannon information analysis in the context of serial descent. A Pleistocene species pump hypothesis is advanced to explain the stenomorphic populations of species of the genus in eastern North America. Key words: Alaska, Anoectangium, British Columbia, bryophyta, China, evolution, hyperoceanic, India, Pottiaceae, Shannon information.

The discovery of the Asian (India, China) species, (Zander & Eckel, 2007), and South America (Cano & Anoectangium sikkimense M. N. Aziz & Vohra, Pottiaceae Jim´enez, 2013). This paper is a synopsis mainly focusing (Zander, 2017a), in Alaska, and the report of A. stracheya- on Anoectangium Schw¨agr. in coastal northwestern North num Mitt., previously known from Asia and eastern North America. Worldwide, about 40 species are accepted. America south through Mexico, Central America, and the Schofield and Crum (1972) have reviewed the exten- Andes, as new to Canada from British Columbia (Zander sive past literature on bryophyte disjunctions and areas & Eckel, 2017), prompted Olivia Lee at the University in hyperoceanic climates, which include, in the North, of British Columbia to segregate 40 specimens of the genus Britain and Norway, the Faroes, the Alps, the Hima- in the herbarium (UBC) as possible specimens of these layas, mountainous Japan and Taiwan, North Pacific species, and to send them to the author for further analysis. North America, and high mountain slopes in the Ha- Lee suggested that one specimen, Schofield 67663,was waiian Islands, and in the South, southern Australia and A. stracheyanum and new to Canada, which proved correct. southern South America. They discussed probabilities The specimen identifications are summarized in the sec- of long-distance dispersal and vicariance explanations tions on representative specimens examined, though for the disjunctions, and concluded that no one hypoth- the work proved to be more complex than expected. esis explains all known data. More recently, Schuster This article provides a chance to gather together in one (1983: 497, 535) mapped and discussed endemic spe- place the new methods used for macroevolutionary cies in the northwestern North American hyperoceanic analysis, such that future Methods sections need only area as exhibiting a Laurasian distribution pattern. reference this paper for basic techniques. Disjunctions involving the northwestern hyperoceanic The present paper helps complete knowledge of the species of Anoectangium are apparently included in his genus studied in its worldwide distribution (Zander, discussion of circum-Laurasian distributions dissected 1993), for Middle America (Zander, 1977), North America by Pleistocene events and survival in refugia.

I am grateful to the Missouri Botanical Garden for continued support for this and similar research. I am also grateful to O. Lee, collections manager at UBC, for bringing significant specimens from British Columbia and Alaska to my attention. L. Briscoe, collections manager at NY, is thanked for loan of specimens. P. M. Eckel kindly provided the illustration. Three anonymous reviewers were also quite helpful. Missouri Botanical Garden, 4344 Shaw Blvd., St. Louis, Missouri 63110, U.S.A. [email protected] Volume 104, Number 2 Zander 325 2019 Macroevolutionary Evaluation with Anoectangium (Pottiaceae) in North America and the Himalayas

There have been five Anoectangium species, A. Anoectangium specimens from the Mitten herbarium aestivum (Hedw.) Mitt., A. euchloron (Schwägr.) Spruce, at NY, including those of A. clarum Mitt., A. crassiner- A. handelii Schiffn., A. sikkimense M. N. Aziz & Vohra, vium Mitt., A. stracheyanum, and A. thomsonii Mitt. A and A. stracheyanum Mitt., reported for North America key (Key 1) to the species of eastern Asia and North (Zander, 1977, 2017a; Zander & Weber, 2005; Zander America using traits as combed from the east Asian & Eckel, 2007, 2017). Anoectangium euchloron was and American literature (particularly Lawton, 1971; treated by Zander and Eckel (2007) as A. aestivum, but Gangulee, 1972; Saito, 1975; Zander, 1977; Eddy, revision by Cano and Jiménez (2013) has demonstrated 1991; Sharp et al., 1994; Li & Iwatsuki, 1997; Li et al., its distinction. Problems in identification of the speci- 2001; Allen, 2002; Aziz & Vohra, 2008; Cano & mens from UBC required examination of types of Asian Jiménez, 2013) is as follows.

KEY 1. KEY TO NORTH AMERICAN AND HIMALAYAN SPECIES OF ANOECTANGIUM WITH DISTINCTIONS AS EXTRACTED FROM THE LITERATURE 1a. Leaves blunt, costa subpercurrent to percurrent. 2a. Leaves 0.7–1.2 mm in length, costa percurrent, gemmae absent ...... A. euchloron (Schw¨agr.) Spruce 2b. Leaves 0.4–0.5 mm in length, costa ending 3 to 4 cells before apex, gemmae often present on rhizoids in leaf axils ...... A. handelii Schiffn. [5 Molendoa antiqua R. H. Zander] 1b. Leaves acute, costa percurrent or more commonly excurrent as a short or stout mucro. 3a. Leaves abruptly widened at base as a broad skirt, basal margins strongly crenulate-denticulate ...... A. sikkimense M. N. Aziz & Vohra 3b. Leaves not or somewhat widened at base, basal margins entire. 4a. Leaves with a weak constriction at top of leaf base, leaves about as wide above as at leaf base ...... A. stracheyanum Mitt. 4b. Leaves without a constriction at top of leaf base, leaves widest at middle or just below. 5a. Distal laminal cells very thick-walled, papillae distinct, separate ...... A. clarum Mitt. 5b. Distal laminal cells usually thin-walled, papillae crowded and somewhat fused. 6a. Leaves 0.7–1.2 mm in length, costa subpercurrent or more commonly ending in a small apiculus ...... A. aestivum (Hedw.) Mitt. 6b. Leaves 1.2–2.5 mm in length, costa commonly excurrent as a large apiculus or mucro ...... A. thomsonii Mitt. [5 A. aestivum (Hedw.) Mitt.]

METHODS extension its roots in intuitive taxonomy, is firmly rooted in scientific theory and philosophy. The most important The aim of macroevolutionary systematics is to de- aspects of these methods are as follows: crease the coding of hidden information (negentropy) about evolution. Its methods are often at odds with those 1. Neo-Husserlian bracketing—In philosophy, this re- of phylogenetics. Classical taxonomy, although in part quires reevaluating a group from scratch, while an intuitive art form based on long familiarity with the using past work as guidelines, not holy writ, in an range of variation in a taxon, is not magic. Special effort to largely eliminate bias, beliefs, and values scientific techniques used in this macroevolutionary (Zander, 2018: 167). Any revision follows this con- study—and doubtless in part unconsciously by all cept to a large extent. This is why two keys are taxonomists—are summarized here, and many are dis- presented here, to emphasize major differences in cussed in greater detail by Zander (2013, 2014, 2016, present and past species concepts. 2018) though scattered in this literature. These methods 2. Inverse problem solving—This is the positing of as well as some new ones are gathered here in simplified several different solutions that may clarify what form as a Methods section. the problem might be, then testing for the most Systematics is occasionally described as a descrip- effective, essentially the inverse problem-solving tive endeavor, lacking the rigor and clearly supported method in physics. A welter of information may be results of experimental science. Yet, systematics is addressed by induction through informed sorting within the purview of “quasi-experimental” science that reveals suggestions of causal connections. (Cook & Campbell, 1979), which details how to use Problems with theory can be addressed by brave naturally occurring control groups to infer valid results invention of new theory. For instance, intellectual to systematic, ecological, and biogeographic analyses. discomfort with cladistic reasoning was relieved by Intuitive methods (Gestalt, omnispection) have been substitution of the dissilient (radiative) genus con- respected because they “work,” providing integrated cept using direct descent; this illuminated the fact and predictive classifications. Below are some of the that dichotomous trees and node-to-node evolution- previously unclarified reasons why they work. The ary continuity did not model evolution, which has reason for the fancy, unfamiliar terminology is to dem- not been obvious for the past three decades. It also onstrate that macroevolutionary systematics, and by substituted explanations based on process for those 326 Annals of the Missouri Botanical Garden

based on pattern. Heuristic methods of cladistics obvious (Zander, 2018: 36). Anoectangium is a are a kind of inverse problem solving, but all morphologically rather isolated genus and is here phylogenetic solutions are couched in terms of an purified into what may be taken to be a closed optimal dichotomous tree. causal group by elimination of A. crassinervium and 3. Abductive sampling—Inferential reasoning from the A. handelii, which belong (the latter as synonym) significance of one fact or facts to the significance of with Molendoa Lindb. Systematics builds on the another fact. This is much like Sherlock Holmes’s work of others by using prior classifications as ever- identification of a particular criminal from partic- better attempts at detailing closed causal groups. ular evidence. Morphological studies are usually 7. Causal asymmetry—Evolution may be modeled as a well sampled, except for very rare taxa for which causal series. Inferring evolutionary order by start- evolutionary trends (particularly the kinds of spe- ing with a putative generalist species and identify- cializations) known for congeners and close rela- ing more specialized species as effects of evolution tives allow evaluation of progenitor-descendant is easier and depends on less information and relationships. Likewise, they may reveal possible informational analysis than hypothesizing ancestral major traits that reflect ancestry, identify good species from often reduced or specialized descen- evolution-based taxonomic traits, and contribute dants (see Thompson et al., 2018). Likewise, in to adaptive explanations, as opposed to minor traits molecular systematics, serial evolution is easily that pop up among many species and reveal little deduced from paraphyly of ancestral molecular about relationships. Fitzhugh (2006, 2008) has races, while descendant-to-ancestor connections discussed abduction in the context of modern are difficult to infer from information lacking mo- systematics. lecular paraphyly. Interpreting patterns as causal 4. Granger causality—When eliminating an aspect of processes through time also addresses all data, not theory would potentially remove meaning (predic- just that used to infer shared ancestry of sister tive value) from a classification (Zander, 2018: groups. 167), the theory is then demonstrated as lean, 8. Parallel evolutionary series—If a species is not causal, and productive of a more complete and clearly a member of a particular genus, i.e., not a better supported classification. One rearranges spe- member of a closed causal group, then an evolution- cies definitions or possible relationships until they based classification may be had by recognition that match or fit best evolutionary and biogeographical the somewhat anomalous species may easily fit theory. Eliminating a taxon from a cladogram does into a taxon transformation series of an evolution- little insult to phylogenetic classification, but arily close group. This proved to be the case in the eliminating a process-based caulistic (nested stem present paper with Anoectangium handelii recog- taxa) relationship reduces information and under- nized as a Molendoa, a genus in which it fit well in standing in the context of theory. the stature gradient among its species. At the trait 5. Two-trait species concept—A minimum of two traits’ level, characters commonly have a distinctive spec- difference is necessary to define a species, because trum of variation that may help clarify them. For one trait may simply be a rogue allele (Zander, instance, the low, simple laminal papillae of the 2018: 169). The 2-trait minimum is most evolution- type of A. clarum could have been simply a reduced arily relevant if the traits of the ancestral species are version of the bifid to multiplex papilla variation of known and can be compared. Most species in larger A. aestivum, but specimens with dense, almost furry genera seem to have accumulated, through adapta- exhibition of simple papillae demonstrated that the tion and drift, a total of three or four important traits trait had its own range of expression as biologically during and after speciation. The exception is for expected. morphologically quite isolated species when con- 9. Dissilient genus concept—A genus may be defined geners are extinct, and traits of progenitors have in an empirical manner as a radiative group con- added up to more than a few traits in the isolated sisting of a central, often polymorphic generalized species. species generating one or more short inferred lin- 6. Closed causal group—Analysis of evolutionary re- eages of more specialized descendant species (Zander, lationships is easier when, within reason, none of 2018: 180). The descendants are more similar to the species studied can be assigned to any other the putative progenitor than they are to each other, known genus. In other words, infrageneric relation- as has been pointed out by others. “Dissilient” ships are more easily established when one is means exploding through radiation that may be working with a group with similar traits undergoing nearly neutral (“nearly” because all traits must short, stepwise transformations during speciation, survive the test of phyletic constraint, including in which case the ancestral species is usually fairly those associated with drift) or adaptive or both. Volume 104, Number 2 Zander 327 2019 Macroevolutionary Evaluation with Anoectangium (Pottiaceae) in North America and the Himalayas

Nearly neutral traits resist (are mostly invisible to) and caveats given by Zander, 2018: 74 and follow- selection pressures and persist across speciation ing pages). The summed bits of each progenitor- events. They provide information on descent with descendant pair or of a lineage may be directly modification. New adaptive traits may become nearly translated to Bayesian posterior probabilities (BPP) neutral in descendants. of the order of ancestor-descendant evolution 10. Macroevolutionary evaluation using an “analytic (Zander, 2018: 173), i.e., the chance of a reversal key”—An indented prose diagram shows the of the order of a taxon in the lineage. Summing the inferred direct evolutionary relationships be- bits of all congeners gives a measure of the coherence tween progenitors and descendants (Zander, of the genus in a closed causal group. The Bayesian 2018: 64). It may be extended diagrammatically support for morphologically based caulograms can into a “caulogram,” or evolutionary tree showing be compared to Bayesian support for molecular serial relationships of stem taxa. Inferring direct cladograms to the extent that the trees are com- descent involves an actual model of evolution. It mensurate, i.e., how the species group, not how is more parsimonious than phylogenetic shared they exhibit serial descent. The higher the number descent from an unknown ancestor, because re- of bits, the greater is the entropy (degree of de- versals of the distinguishing traits of the pre- coding of hidden evolutionary information) in a sumptive shared ancestor are not taken into classification. — account in cladistic parsimony estimation; see 12. Names of extant species as progenitors Of course caveats about simplicity by Coveney and High- the ancestors, as individuals and populations, of an field (1990: 267). extant species are no longer alive (except as long- lived clones perhaps), but if the past, long-dead In direct serial evolution, reversals are minimized or individuals are inferred to have about the same eliminated. Each of the often several short lineages of morphology as an extant taxon, that taxon name successively more specialized descendants from a gen- should be used, at least as an effective locum tenens eralized progenitor builds on a set of homologous serially pending new discoveries. Thus, through a kind of nested traits. Very large genera have not yet been sub- evolutionary uniformitarianism based on an expec- jected to nested serial analysis, and one might expect new tation of age-long stasis of species being common, macroevolutionary features associated with massive ad- inferred evolutionary continuity may be consid- aptational sorting. The primary macroevolutionary eval- ered from taxon to taxon, not node to node as in uation is simply a spreadsheet detailing species ordered cladistics. by their nested traits (Table 1). The closest ancestor- descendant difference is two traits because a minimum of two traits is required to distinguish a species. In practice, RESULTS however, descendant lineages may be two to four species or occasionally more in length (e.g., see the caulograms of TAXONOMY Zander, 2014, and Zander, 2018: 70; also Fig. 1). This Study of the UBC material and the past analysis of may include new generalized species that establish their specimens from elsewhere in the world (see Zander, own different dissilient genera. Inferred “missing links” 1977, 1993: 138 for lists of herbaria) found that leaf are allowable when direct descent exceeds expected length, basal marginal crenulation, and relative costa gradual accumulation of traits along lineages (punctua- length of the leaf were not particularly good traits to tional or anagenetic), which is observed to be generally distinguish species because of considerable overlap and three to four new traits per speciation event in past variation. Leaf shape, however, is fairly constant and macroevolutionary evaluations. correlates with apiculus length and other traits as noted 11. Sequential Bayes analysis—Simply speaking, ho- in Key 2. Clumps of Anoectangium species are steno- mology assessment and closing the causal group morphic within the clump (large tuft, small cushion), but these are often different from other clumps in remove “noise” by synonymizing redundant traits morphology, particularly in plant size. Although female and leave only evolutionarily significant differ- gametangia are common in Anoectangium specimens ences. Traits distinguishing a descendant species examined, no sporophytes were found in the UBC from its ancestral species may each be assigned one material. The species in northwestern North America bit of Shannon information. A bit consists of a apparently only reproduce asexually. It is then hypoth- morphological, ecological, or geographic trait that esized that sexual mixing is not the source of the rather aids in determining order of nesting of taxa. Mo- free combinations of traits important in previous studies lecular traits may be used if they, too, can contribute but that the hyperoceanic climate, multiplicity of hab- to understanding serial evolution (see discussion itats in British Columbia and southeast Alaska, and 328 Annals of the Missouri Botanical Garden

Table 1. Macroevolutionary analysis of Anoectangium Schw¨agr. species in the New World and Himalayas showing traits of progenitor and inferred descendant species, each trait awarded one informational bit, these converted to Bayesian posterior probabilities (BPP) of support for order of descent. Vertical lines imply speciation from an ancestral species to the left and above, and degree of indentation shows joint descent. Each descendant shares homologous traits with its progenitor, but differs by inferred evolutionary new traits given in the fourth column.

Second Ancestral traits and new Support for order Ancestor Descendant descendant traits of descendants of descent A. aestivum leaves equal-sized, elliptic to lanceolate, . 0.6 mm, acute, apiculate to short-mucronate, base rarely serrulate-crenulate; distal laminal cells homogeneous, unistratose, thin- to thick-walled, moderately convex, rounded-quadrate in section, papillae massive, usually fused, ca. 3 to 4 centered over each lumen; gemmae absent A. euchloron rosulate, leaves long-elliptic to ligulate, 5 bits, 0.97 BPP apex broadly acute to rounded; distal laminal cells low spiculose-papillose, walls nearly circular in section A. clarum leaves long-lanceolate to long-triangular, distal 4 bits, 0.94 BPP laminal cells very thick-walled, lumens usually oval; laminal papillae small, simple, ca. 4 to 5 evenly scattered across each lumen A. incrassatum leaves short-elliptic, apiculus short or absent, 4 bits, 0.94 BPP medial cell lumens rounded-rectangular, distal marginal cells elongate A. stracheyanum leaves long-lanceolate to long-ligulate, length 3 bits, 0.80 BPP 7 to 10 times greatest width above leaf base, often constricted just above the weakly sheathing leaf base A. sikkimense leaves long-triangular, base abruptly widened 3 bits, 0.89 BPP as a short-sheathing skirt about 1.3 times as broad as at midleaf, leaf base often serrulate-crenulate on margins

perhaps lack of competition on wet rock ledges have is a taxonomically difficult genus but not impossible. preserved many combinations of possible traits. A revised key (Key 2) to these species of Anoectangium Given that species of similar moist habitats (Ardeuma as they appear in the Himalayas (Aziz & Vohra, 2008), recurvirostrum (Hedw.) R. H. Zander & Hedd., Gymnos- North America, and generally in the New World, building tomum aeruginosum Sm., species of Philonotis Brid. and the on the studies of Cano and Jim´enez (2013), Zander (1977), like) are also rather variable in morphology, Anoectangium and Zander and Eckel (2017), follows.

KEY 2. REVISED KEY TO SPECIES OF ANOECTANGIUM IN NORTH AMERICA AND THE HIMALAYAS 1a. Distal laminal cells very thick-walled and lumens oval or rounded-rectangular, laminal papillae small, simple, scattered over the lumens, usually not or little obscuring the cells. 2a. Leaves long-lanceolate to long-triangular, occasionally long-elliptic, mucro absent to stout, distal marginal cells similar to the medial cells, medial cell lumens oval ...... 2.A. clarum Mitt. 2b. Leaves short-elliptic, short-apiculate or apiculus absent, distal marginal cells elongate in one row, medial cell lumens rounded-rectangular ...... 4. A. incrassatum Broth. 1b. Distal laminal cells moderately thick-walled and lumens rounded-quadrate, or very thickened only near costa or thin- walled, laminal papillae large, low-spiculose or dome-shaped and often fused into a blunt capitulum, centered over the lumens, obscuring the cells. 3a. Leaves long-triangular, finely pointed, base abruptly widened as a short-sheathing skirt, or occasionally long- sheathing, 1.3–1.5 times as broad as at midleaf, skirt usually filled with short-rectangular, incrassate cells, leaf base commonly serrulate on margins ...... 5.A. sikkimense M. N. Aziz & Vohra Volume 104, Number 2 Zander 329 2019 Macroevolutionary Evaluation with Anoectangium (Pottiaceae) in North America and the Himalayas

Figure 1. Caulogram of Anoectangium Schw¨agr. species in North America and the Himalayas. Number of bits are the number of traits distinguishing a species from its direct progenitor. Bayesian posterior probability (BPP) of being a descendant is calculated directly from the number of Shannon informational bits. Total bits supporting the coherency of the genus are 19. The bits of each of the two 2-taxon divergent lineages may be added to gauge support for each, which is quite high. Anoectangium aestivum (Hedw.) Mitt. is here considered the supergenerative central taxon of the dissilient genus, being generalist, biotype-rich, widespread in many habitats, and most similar to the outgroup Streblotrichum P. Beauv. Anoectangium stracheyanum Mitt. and A. clarum Mitt. are each also progenitors of descendant species, while A. euchloron (Schw¨agr.) Spruce is apparently evolutionarily isolated. 330 Annals of the Missouri Botanical Garden

3b. Leaves elliptic to long-lanceolate, blunt to narrowly acute, base as wide as at midleaf or less, if wider then weakly so, not or weakly sheathing, leaf base with cells short-rectangular and incrassate only medially, basal margins rarely serrulate. 4a. Leaves long-lanceolate to long-ligulate, rarely long-triangular, leaf length (5 to)7 to 10 times greatest width, often constricted just above the weakly sheathing leaf base; often with a stout bistratose cylindric mucro of 5 to 10(to 30) smooth cells ...... 6. A. stracheyanum Mitt. 4b. Leaves ligulate to elliptic or lanceolate, leaf length less than 7 times greatest width, without a constriction; often with a small apiculus or unistratose flat or channeled mucro of 3 to 8(to 20) smooth cells. 5a. Plants often rosulate; leaves long-elliptic to ligulate, leaves widest at midleaf or near apex, margins usually nearly straight and parallel, apex usually broadly acute to rounded, apiculus small; distal laminal cells spiculose-papillose, walls strongly bulging and nearly circular in section with internal walls less than half the extent of the external walls, cells sometimes crumpled into transverse rows ...... 3.A. euchloron (Schw¨agr.) Spruce 5b. Plants with leaves of similar length all along stem; leaves elliptic to lanceolate, leaves widest at midleaf or just below, margins arcing, apex acute, apiculus small to enlarged as a mucro; distal laminal cells with dome-shaped, multiplex-fused papillae, moderately bulging and rounded-quadrate in section with internal walls more than half the extent of the exposed walls, occasionally nearly flat superficially, cells never crumpled ...... 1.A. aestivum (Hedw.) Mitt.

1. Anoectangium aestivum (Hedw.) Mitt., J. Linn. 2012 as A. aestivum, has all traits of that species except Soc. Bot. 12: 175. 1869. Basionym: Gymnostomum that the section of the lamina reveals the string-of-beads aestivum Hedw., Spec. Musc. 32. 1801. TYPE: bulging cells with small interconnections typical of Hedwig, Sp. Musc. Frond., tab. II fig. 4-7. 1801 A. euchloron. This is an example of the multiple biotype (lectotype, designated by Geissler [1985]). feature of the species A. aestivum, which supports recognition as progenitor of the extant lineage. Anoectangium thomsonii Mitt., J. Proc. Linn. Soc., Bot., Suppl. 1: 31. 1859, syn. nov. TYPE: [India], Simla, 7000 ft., Variability may enhance fitness. According to T. Thomson 156 (lectotype, designated by Li & Iwatsuki Ferrière and Fox (1995), the chaotic distancing of bio- [1997: 249], NY!). types as a kind of emergent speciation enhances the Anoectangium aestivum is worldwide in distribution survivability of the species metapopulation, as studied in temperate and montane areas. Examination of spec- by Allen et al. (1993), who interpreted biotype increase “ ” imens of A. aestivum from Europe and Asia found that as evolutionary noise. The large-stature gradient in most specimens had distinctly bulging laminal cells, Anoectangium aestivum may be investigated with mo- an apiculus or short and sharp mucro, and leaves lecular analysis. Finding certain morphotypes having that graded from short-lanceolate in small plants to multiple molecular races as descendant lines implies a long-ovate in large forms. None of the specimens from potential status as subspecies or variety because mere northwest North America had sporophytes, although phenotypic variation should possess the same races as perichaetial buds were common. One specimen from their normative morphology. This would require exten- Alaska (Worley 8784, MO) had black leaves fragile in sive sampling. pieces except for the most terminal and may prove a Some of the syntypes of Anoectangium thomsonii sent good variety. from NY cited by Mitten (1859) are A. stracheyanum, Some subtropical and tropical specimens of Anoec- and two from India, Strachey & Winterbottom s.n. and tangium aestivum s. str. intergrade with the southern Royle s.n., are A. sikkimense, but one specimen from species A. euchloron. For instance, Morley & Whitefoord India, Thomson 156, which fits the protologue and was 731 (MO), from Jamaica, has leaves broadest in the previously selected as lectotype by Li and Iwatsuki middle or beyond and blunt apices, but is not rosulate, (1997: 249), is a large form of A. aestivum that matches the leaves are not twisted around the stem when dry, current usage for A. thomsonii. I here conclude that laminal cell sections are rounded-quadrate and interior A. thomsonii is simply a large form of A. aestivum of local walls are about the same breadth or larger than the montane or hyperoceanic distribution. Typical of a large superficial walls, and the laminal papillae are low, form of A. aestivum is a specimen from British Columbia, crowded, and hemispheric. Another intergrading spec- Schofield & Boas 21574 (MO), with leaves ca. 1.8–2mm imen is Brinson s.n. (MO), from 1979 in El Salvador, in length, 4–5:1 in dimension, with perichaetiate buds. which is also not rosulate (the leaves are evenly spaced), A thorough revision is needed, but the distribution of the leaves are not twisted when dry but are widest at A. aestivum as reported in the Flora of North America midleaf or beyond, the apex is rounded, the laminal cells (Zander & Eckel, 2007) is correct. in section have round lumens, and papillae are short- Representative specimens examined. CANADA. British spiculate; thus, it is A. euchloron. A specimen from Columbia: toward head of Kingcome Inlet, N side, humid Colombia, Cleef 7384 (MO), identified by M. J. Cano in cliff face near waterfall, Schofield 41324 (UBC); Jervis Inlet, Volume 104, Number 2 Zander 331 2019 Macroevolutionary Evaluation with Anoectangium (Pottiaceae) in North America and the Himalayas head of Queen’s Reach, mouth of Lausmann Creek, humid cliff identical in sharp leaf apex with authentic Himalayan shelf, Schofield 38293 (UBC); Howe Sound, Shannon Falls, material, while the specimens from Japan, including shaded humid base of boulder, Schofield 82733 (UBC); Prince Inoue 427 (MO), have elliptical leaves with large, Rupert, Kalen Island, Hays Mtn., Schofield & Boas 21574 (MO); Vancouver Island, near Vancouver, Wigwam Creek, crowded but evenly spaced simple to occasionally canyon walls, Schofield 20530 (UBC); Haida Gwaii, Moresby short-bifid papillae, appearing like a coat of short Island, S side of mouth of Tasu Sound, shaded cliff crevices, fur and often obscuring the lumens. Anoectangium Schofield 37475 (UBC); Graham Island, McClinton Bay, Mas- euchloron is similar to A. clarum and may have evenly set Inlet, damp cliff crevice, Schofield 35244 (UBC); NE side of Hibben Island, NW coast of Moresby Island, Schofield 15358 scattered, small, simple distal laminal papillae as is (UBC); Chasti Island, damp cliff, Schofield 18783 (UBC). the case with A. clarum, but the laminal cells are CHINA. Yunnan: Baoshan, Teng-Chong Co., Jiang-Zu Vil- bulging in the former and have thin or only moderately lage, Gao-Li-Gong-Shan NNR, Lin-Jia-Pu Forest Station, W. Z. thickened walls, e.g., Fuentes & Aldana 6464 (MO), Ma 14-5524 (MO). COLOMBIA. Boyacá: Duitama, Páramo from Bolivia. A specimen from Hawaii, Bartram 292 de La Rusia, Cleef 7384 (MO). INDIA. Kumaon: Devi-dhura, 7000 ft., s.d., Strachey & Winterbottom 19 (BM). Simla: 7000 (FH!), the type of A. haleakalae var. laxulum E. B. ft., T. Thomson 156 (NY—lectotype of taxonomic synonym A. Bartram, is also similar to A. clarum except for some- thomsonii). JAMAICA. Portland: Blue Mtns., Nannypot Is- what larger papillae, these likewise simple, hemi- land to Kill Head, woodland, ca. 2600 ft., Morley & Whitefoord spheric, and scattered. If after additional study 731 (MO). U.S.A. Alaska: Sitka Co., Alexander Archipelago, Baranof Island, Thimbleberry Lake Trail, Schofield 122951 many descendant species are discovered for A. clarum, (UBC); Sawmill Creek, below Blue Lake, Worley & Hamilton supported perhaps by demonstrable molecular race 9456 (UBC); S of Ketchikan, roadside waterfall, Worley & paraphyly (Zander, 2018: 18, 74), this would imply Thorpe 2849 (UBC); San Juan Bautista Island, SW of Craig, that it is the supergenerative progenitor for a new Worley & Hamilton 7250 (UBC); Mithof Island, 26 mi. S of dissilient genus. Petersburg, Worley 8784 (MO). Washington: Snohomish Co., Granite Falls, wet cliff, Schofield et al. 21892 (UBC). Representative specimens examined. CHINA. Henan: Songxian Co., Luo 443 (MO). Zhejiang: Mt. West Tian-Mu, 2. Anoectangium clarum Mitt., J. Proc. Linn. Soc., old temple, R.-L. Hu 54 (MO). INDIA. Maharashtra: Ling- Bot., Suppl. 1: 31. 1859. TYPE: India. Sikkim: malla, Sedgwick 4659 (MO). Punjab: Kangra, Kulu, Bushreo Himalaya, Lachsong, 15,000 ft., s.d., Hooker 203 La, Koelz 7512 (MO). Sikkim: Lachoong, 15,000 ft., J. D. Hooker 203 (NY—lectotype, MO). Uttaranchal [Uttarak- (isotype, MO!; lectotype, designated by Li & hand]: Nainital Distr., Bareilly College, Saxena & Kumar Iwatsuki [1997: 246], NY!). 20911000268D (MO). JAPAN. Miyazaki: Kaeda Valley, near Mt. Boroishi, 1977, Inoue s.n. (Bryophyta Selecta Exsiccata Anoectangium clarum has been known from China 427) (MO). PHILIPPINES. Luzon: Bued Cañon, Bartlett and India and here is found new to Japan and the 13394 (DUKE, MO). Philippines. It is similar to large forms of A. aestivum in leaves to 2.5 mm. It differs in the long-lanceolate to 3. Anoectangium euchloron (Schwägr.) Spruce, Cat. long-triangular leaves with a stout mucro, and evenly Musc. 4. 1867. Gymnostomum euchloron Schwägr., spaced, scattered, punctiform laminal papillae. The trait Spec. Muse. Suppl. 2(2): 83. 1827. TYPE: West combinations characteristic of A. clarum were not found Indies, Martinique, s.d., Sieber s.n. (G!). in the UBC material from British Columbia and Alaska, The distribution of Anoectangium euchloron is es- or indeed among any of the New World material seen. sentially the same as that reported by Cano and Jiménez Many specimens of A. aestivum have incrassate laminal (2013), being known from North America, Central cells, but the papillae are always enlarged, centered America, South America, and across Asia in warm- over and obscuring the lumens. The treatment of Anoec- temperate and tropical areas. Material with the strap- tangium in China by Li et al. (2001) presented A. clarum shaped leaves and nearly circular cross-section of in a different light than exhibited by the type at NY. laminal cells of A. euchloron was not found in the This species was originally named “clarum” because UBC material. The degree of laminal cell wall protrusion the ovate laminal cell lumens were clearly visible under is used by Cano and Jiménez (2013) in part to distin- the tiny punctate papillae. Although the key of Li et al. guish A. aestivum and A. euchloron. Degree of cell (2001) reflected this, their description of A. clarum, bulging can be best examined with leaf sections, but illustration of it, and annotations on several specimens an easy method of determining the bulging of the cells is from NY all indicate that thick cell walls with large, by optical section of the edge of a leaf about midleaf. rounded papillae were included in their concept. Previous treatments of Anoectangium euchloron by Anoectangium clarum was included in the present Zander (1977) and Zander and Eckel (2007, 2017) study because of the traits it shares with A. euchloron treated this species as a synonym of A. aestivum. and A. incrassatum. Anoectangium clarum is doubtless Cano and Jiménez (2013), however, distinguished A. widespread in Asia. A specimen from the Philippines, euchloron from A. aestivum by the bulging laminal cells Bartlett 13394 (DUKE!), as A. subclarum, is nearly of the former versus the flat surfaces of the laminal cells 332 Annals of the Missouri Botanical Garden

of the latter. Specimens of A. aestivum with very thin cell A. aestivum by Allen (2002) for Central America, though walls may have rather flat cell superficial walls in some he recognized the difference between the two variants; plants of a collection, while the basal laminal cells A. euchloron is confirmed here for El Salvador and always have flat surfaces. The image of the leaf section Panama. of A. aestivum in the synopsis by Cano and Jiménez Representative specimens examined. BOLIVIA. La Paz: (2013), on the other hand, shows weakly bulging and Franz Tamayo, R´ıo Machariapo, 12 km al N-NE de Apolo, clearly convex cell walls, which are not “plane” as Fuentes & Alana 6464 (MO). EL SALVADOR. Cuscatlán: stated in their key although definitely not strongly Laguna Las Ninfas, thin soil over rocks, 1970, Brinson s.n. — bulging. The key given here clarifies the cautions (MO). MARTINIQUE. s. loc., s.d., F. W. Sieber s.n. (G holotype of Gymnostomum euchloron); Duss 365 (NY). MEXICO. Mor- offered by Zander and Eckel (2017) regarding apparent elos: near Cuernavaca, damp cliffs, 5000 ft., Pringle, Plantae intergradation in northern North America between A. Mexicanae 543 (MO). PANAMA. Chiriqu´ı: 2 mi. S of Boquete, euchloron and A. aestivum, i.e., there is little intergra- rotting log, Crosby 3969 (MO). U.S.A. Arizona: Santa Cruz dation based on the present reevaluation of the traits. Co., Patagonia Mtns., Flux Canyon, 1524 m, Bartram 971 Some specimens (e.g., Schofield 15358 and Schofield (DUKE, FH). 41324, UBC) from British Columbia have distal laminal 4. Anoectangium incrassatum Broth., Bot. Tidsskr. cells distinctly rounded in section, with internal walls 36: 279. 1919. TYPE: Jamaica, Børgesen s.n., about half the breadth of a superficial wall, as is the case 1906 (H not seen). Figure 2. with the southern species A. euchloron; these specimens, however, have the lanceolate leaves and rounded, A survey of slides reviewed by Zander (1977) and an centrally crowded papillae characteristic of A. aestivum examination of collections at MO revealed that an (see Key 2). apparent descendant species of Anoectangium clarum Some specimens of Anoectangium euchloron have is present in the West Indies as A. incrassatum; see lists laminal cells irregularly bulging on both sides of the of representative specimens examined. Anoectangium leaf, e.g., Lewis 79-503 (MO!), from Bolivia, or even in incrassatum is known from Cuba, Dominican Republic, regular, transverse rows, e.g., Duss 365 (NY), from Haiti, Jamaica, and Puerto Rico and is discussed and Martinique, illustrated by Zander (1976). The type of cursorily illustrated by Crum and Steere (1957: 460) A. euchloron at G has leaves 0.8–1 mm in length, long- with additional information by Crum and Bartram ligulate, with a percurrent costa, 4-celled apiculus, (1958). Anoectangium incrassatum is included here broadly acute apex, no basal constriction, smooth basal as similar to A. euchloron in the short-elliptic leaves margins, thickened laminal cells and short-spiculose, and subtropical distribution. It is apparently geograph- crowded papillae. The apex was definitely acute, not ically disjunctive to Japan. A specimen from Japan, blunt, as is also shown in plate 177 of Schwägrichen Schofield 51907 (MO), is a robust version of A. incras- (1827), although A. euchloron commonly has rounded satum with longer, more apiculate leaves, more acute leaf apices. Although also from Martinique, the type of apex, and more crowded laminal papillae, but matches A. euchloron does not have laminal cells bulging in in thick-walled laminal cells with squared lumens, transverse rows, like that of Duss 365 (NY); instead the elongate marginal cells, and simple laminal papillae. cells bulge out irregularly and individually. A similar The species is new to Japan. In the Americas, the condition occurs in A. wilmsianum (Müll. Hal.) Paris twisting of the leaves is not particularly distinctive as of South Africa (Zander, 1993: 138), which may be it varies through states much the same as does that of synonymous. A. euchloron and probably is associated with a constraint A specimen from Arizona, Bartram 971 (DUKE, FH), on flexion due to the short leaves. reported by Bartram (1927), is indeed Anoectangium Anoectangium incrassatum may be interpreted as euchloron, apparently the only known collection for the an evolutionarily much reduced species, with short- United States; Breen (1963) and Reese (1984) did not elliptical leaves about 0.6–0.8 mm long, rounded include the genus in their southern U.S. identification apex, and short or absent apiculus. Given the scat- manuals. The species is distributed in Mexico and the tered geographic range, A. clarum and A. incrassa- West Indies, and is widely distributed in South America tum, as part of a lineage, are rather ancient. The (Cano & Jiménez, 2013). Representative specimens presence of a species of Asian lineage in the West seen are commonly short stemmed, rosulate or nearly Indies presents an interesting phytogeographic so, leaves when dry are conduplicate-incurved, and in a problem, possibly involving an ancient Tethyan sub- weakly twisted Z form (as seen from the side, or to the tropical distribution (Schuster, 1983: 491; Rodr´ıguez- left, or counterclockwise going up when looking down Sánchez & Arroyo, 2008; Wen & Ickert-Bond, 2009). from the top of the plant, the twist of a standard screw), This curious disjunction involves the broadleaf ever- or occasionally in an S form (the opposite twist) or green sclerophyllous vegetation bordering an Early not at all. Anoectangium euchloron was included in Tertiary sea stretching from southwestern North Volume 104, Number 2 Zander 333 2019 Macroevolutionary Evaluation with Anoectangium (Pottiaceae) in North America and the Himalayas

Figure 2. Anoectangium incrassatum Broth. —A. Moist habit. —B. Dry habit. —C. Perigonium. —D. Perichaetium with emergent calyptra. —E. Stem section. —F. Two leaves. —G. Leaf apex. —H. Leaf base. —I. Laminal papillae. —J. Leaf section near base. —K. Leaf section near apex. From Ekman 1918 (MO). Scale bars: a 5 2mm(A–D); b 5 3 mm (F); c 5 25 mm (E, G, H, J, K); d 5 10 mm (I).

America to south-central Asia, and is often cited as States, northwestern South America, eastern Asia, determining disjunctions between Mediterranean cli- and Indonesia. Given that bryophyte species may be mate areas of California and southern Europe (but see in morphological stasis for millions of years, and that Shaw et al., 2003). The same explanation may suffice many species have nearly worldwide ranges, no spe- for the distribution of A. incrassatum–A. clarum and, cies should be considered too geographically isolated for instance, that of Luisierella barbula (Schw¨agr.) for allopatric speciation from a similar progenitor that Steere, found in the West Indies, southern United is often extant. 334 Annals of the Missouri Botanical Garden

Representative specimens examined. HAITI. Ouest: Mas- selected a specimen from Kumaon, India, Strachey & sif de la Selle, Furcy, gorge of Riviere Tebaud, shady rocks, Winterbottom 19 (NY), as lectotype. The protologue, – 1000 1100 m, Ekman 1918 (MO). JAMAICA. Blue Moun- however, cites “Strachey et Winterbottom” but does not tains: Abbey Green, 5000 ft., 1928, Orcutt s.n. (DUKE, MICH, MO). JAPAN. Nagano Pref.: Kinofuku-shima, Schofield give a collection number. Two specimens of Strachey & 51907 (MO). Winterbottom 19 were made available by BM, and both were of small plants that were taxonomically A. aesti- 5. Anoectangium sikkimense M. N. Aziz & Vohra, vum. The second syntype of A. stracheyanum (from – Bull. Bot. Surv. India 30: 187, f. 1 14. 1988 Simla, India, 7000 ft., Thomson 207) was also commu- [1990]. TYPE: India, Sikkim, Sinchul, Kurz nicated by BM. This proved to be A. stracheyanum 2095 (holotype, CAL not seen). taxonomically. Mitten (1859) also cited Wilson (1857: Generally, Anoectangium species with large leaves 326) in the protologue, who cited only Thomson 207.It have a longer apiculus, grading into a thick, sharp is evident that the lectotype selection of Li and Iwatsuki mucro. This is particularly true for A. sikkimense, for (1997) is to be superseded (Turland et al., 2018: Art. which the long, tapering, narrowly triangular leaf shape 10.2, 19.9) because the lectotype at NY doubtless does enhances the apparent length of the often cylindric not match the description or citation in the protologue. mucro, which is composed of several cells, although Therefore, Thomson 207 (BM) is the proper lectotype some small specimens have only a sharp but short and is designated as such here, which preserves com- apiculus (e.g., Schofield 38343, UBC). Anoectangium mon usage of the name. stracheyanum often has a similar cylindric mucro (Scho- Some of the specimens from the same locale cited by field 34006, UBC) but lacks the broadened flare of the Mitten (1859) as Anoectangium thomsonii, e.g., Hooker leaf base. 197, 200, 201, and Thomson 153 and 199, are A. Anoectangium sikkimense was recently reported as stracheyanum at NY. These indeed have a constriction new to North America from Alaska (Zander, 2017a), and just above the leaf base, but the constriction is absent or additional records are given here for British Columbia. not evident in all leaves, many of which have no Specimens from northwestern North America with the constriction. The constriction consists of one or two long-triangular leaf shape coupled with flaring base fewer cell rows across the leaf and a somewhat stronger keeling just above the leaf base, at least on a microscope (e.g., 1975, H¨ubschmann s.n., UBC) of A. sikkimense slide. This, too, is a poor taxonomic trait, though better lacked marginal crenulations along leaf bases, or had than that of basal marginal crenulation. only hints of such, although some specimens of A. When clumps of Anoectangium stracheyanum or aestivum had basal marginal crenulations. It is also A. sikkimense grow under harsh conditions, the leaves true that those Himalayan syntypes of A. thomsonii that are shorter, and important distinguishing traits—constriction proved to be A. sikkimense also largely lacked basal of leaf above base, and broad skirt of leaf base, marginal laminal crenulations, a trait that is not par- respectively—are less fully expressed. This is expected ticularly diagnostic. variation in moss species characteristic of wet habitats, Representative specimens examined. CANADA. British and some collections labeled A. aestivum may actually be Columbia: Toba Inlet, N shore, E of Brahm Bay, damp cliff, one of the other two species with reduced morphology. Schofield 38343 (UBC); Vancouver Island, Golden River, 1975, H¨ubschmann s.n. (UBC); mtn. just S of Harrison Lake, Anoectangium stracheyanum is present in the West Skeena River, Schofield 21276 (UBC); Haida Gwaii, Moresby Indies, having been collected by G. L. Smith in 1979 Island, Takakia Lake, S of Moresby Logging Camp, Schofield (see Representative specimens examined). 24888 (UBC). CHINA. Henan: Lanchuan Co., D.-Z. Tu 3102 Amphidium lapponicum (Hedw.) Schimp. (Orthotri- (MO). INDIA. Kumaon: Dwali, s.d., Strachey & Winterbottom chaceae) is superficially quite similar to Anoectangium s.n. (NY); Northwest Himalaya, s.d., Royle s.n. (NY). Both India specimens are syntypes of A. thomsonii excluded from A. stracheyanum in the lanceolate, apiculate leaves and aestivum as circumscribed above. U.S.A. Alaska: Shungnak large, crowded laminal papillae, but differs in the auto- Region, Mauneluk River region, just NE of base of Mt. 2820, icous inflorescence, stem central strand absent, and Worthwhile Cliff, Lewis 483 (NY). somewhat larger leaves at 2.5–3 mm in length, with basal cells filling the elliptical leaf base, thin-walled, 6. Anoectangium stracheyanum Mitt., J. Proc. Linn. with transverse walls often somewhat thickened. Amphi- Soc., Bot., Suppl. 1: 31. 1859. TYPE: India. Simla, dium mougeotii (Bruch & Schimp.) Schimp. is imme- northwestern Himalaya, 7000 ft., Thomson 207 diately distinguished by the minuscule elliptical distal (lectotype, redesignated here, BM!). laminal papillae arranged in longitudinal rows. One Two syntypes of Anoectangium stracheyanum were must be careful with this trait in that Amphidium requested for loan from the Mitten herbarium at NY, but lapponicum also has similar elliptical papillae on the L. Briscoe (pers. comm.) reported that these two spec- basal cells, but its distal laminal cells are large, simple imens are at present unavailable. Li and Iwatsuki (1997) to warty, covering the rather angular lumens. Volume 104, Number 2 Zander 335 2019 Macroevolutionary Evaluation with Anoectangium (Pottiaceae) in North America and the Himalayas

Representative specimens examined. CANADA. British series, and M. antiqua fits handsomely into it. The Columbia: Elaho Area, Sims Creek Valley, Peaches & Creme evolutionary formula for inferred serial relationships Waterfall, Schofield 118626 (UBC); Vancouver Island, Brooks (progenitors are boldfaced) including the most well- Peninsula, Doom Mt., cirque face above “throne,” Schofield 82520 (UBC); North Vancouver, Lynn Creek Canyon, Schofield known species is Molendoa hornschuchiana (Hook.) 67663 (NY); Revelstoke Area, S of Revelstoke, Blanket Creek Lindb. ex Limpr. . (M. sendtneriana (Bruch & Prov. Park, Sutherland Falls, Vitt 22396 (UBC); Haida Gwaii, Schimp.) Limpr. . (M. antiqua, M. ogalalensis (G. L. SE side of West Graham Island, Goose Cove, Athlow Bay, near Merr.) R. H. Zander, M. peruviana, M. schliephackei seastacks, Schofield 34006 (UBC); Moresby Island, Goski Bay, Schofield 31615 (UBC). CHINA. Hubei: Shennongjia Forest (Limpr.) R. H. Zander)). Following the idea of looking Distr., S end of Loyang River gorge, near Pingqian, 1980 Sino- for parallel evolutionary series when trying to place a Amer. Exped. 1376 (MO). Shanxi: Quinling Co., H. Quan 506 species with reduced number and expression of critical (MO). COLOMBIA. Meta: Cerro Nevado del Sumapaz, Que- identification traits, a similar reduction series may be brada El Buque, Cleef 7791 (MO). DOMINICAN REPUBLIC. found in another pottiaceous genus, Gymnostomum Cordillera Central: summit of Alto de la Bandera, 2800 m, . Smith 10322 (MO). INDIA. Kumaon: Dwali, 9000 ft., s.d., Nees & Hornsch.: G. aeruginosum (G. calcareum Strachey & Winterbottom s.n. (NY). All cited India specimens Nees & Hornsch. (G. viridulum Brid., G. mosis are syntypes of A. thomsonii. Sikkim: Lachen, 9000 ft., Hooker (Lorentz) Jur. & Milde)), in which the most reduced 201 (NY); Wallanchoon, 12,000 ft., Hooker 197 (NY); Simla, species are gemmiferous and otherwise sterile. Hattu, 10,000 ft., Thomson 199 (NY); Simla, 7000 ft., Thomson 207 (BM), Thomson 153, 200 (NY). Representative specimens examined. CHINA. Tibet: occid. reg. temp., T. Thomson 127 (NY—holotype of replaced 7. Molendoa antiqua R. H. Zander, nom. nov., non name Anoectangium crassinervium). TURKEY. Western Molendoa crassinervis Broth., Bull. Bernice P. Kurdistan: Cataonia, Handel-Mazzetti 2024 (FH—holotype Bishop Mus. 40: 11. 1927. Anoectangium crassi- of synonym Anoectangium handelii). U.S.A. Colorado: Boulder Co., Hall Ranch Open Space, Weber, Wittmann & nervium Mitt., J. Proc. Linn. Soc., Bot., Suppl. 1: Lehr B-114035 (COLO, MO); Larimer Co., Buckhorn Creek, 31. 1859. TYPE: China, Tibet, occid. reg. temp., NNW of Masonville, Hermann 27016 (COLO). Thomson 127 (holotype, NY!). EVOLUTIONARY ANALYSIS Anoectangium handelii Schiffn., Ann. K. K. Naturhist. Hof- mus. 27: 490. 1913, syn. nov. TYPE: Turkey. Western A macroevolutionary analysis of the species of Anoec- Kurdistan, Cataonia, Handel-Mazzetti 2024 (holotype, tangium in the New World using the techniques of the FH!). “analytic key” (Zander, 2013, 2016, 2018), as explained The holotype of Anoectangium crassinervium at NY in short by Zander (2017b), is shown in Table 1. The was examined. This material is a species of Molendoa by analysis is essential in providing evolution-based taxo- the lateral gametoecia, blunt leaves, and low scabriform nomic distinctions between the species of the dissilient or absent papillae. It is the same as what has been (radiative) genus, in addition to detailing their relation- known as A. handelii (Zander & Weber, 2005) by the ships. The study was critical for accurate identification of ovate leaves with subpercurrent costa, and no apiculus, the northwestern American specimens of Anoectangium. and somewhat ragged and crenulate leaf margins. It Alternatively, a classical key would have been problematic lacks the occasional bistratose leaf margin and gemmae in that it usually eliminates the most specialized species of North American material, but this is deemed of little first and forces inferred relationships into a dichotomous consequence in this genus of extremely variable mor- nesting. Formal descriptions themselves give all traits, phology. The epithet crassinervis was previously pub- including minor traits that accumulate as scattered in the lished in Molendoa, hence the new name is proposed genus and related genera. Even when important traits are here. italicized in a description, the way homologous traits are Although large plants of Molendoa species have two shared serially is not demonstrated. costal stereid bands, the single, central stereid band of The evolutionary formula (Zander, 2013: 57, 2018: 6, M. antiqua is like that characteristic of Anoectangium, 46, 81) for the genus Anoectangium in the New World and but this is almost certainly due to the morphological Himalayas is A. aestivum ((A. euchloron)(A. clarum . reduction in costal complexity accompanying general A. incrassatum)(A. stracheyanum . A. sikkimense)); reduction in size. In addition, the thickened cell walls compare Figure 1. The chosen outgroup is Streblotrichum along the leaf margin and low, flattened laminal papillae P. Beauv., from which Anoectangium differs in the are characters that delimit Molendoa. Plants of A. advanced traits of perichaetia lateral on short branches, handelii are quite similar to those of M. peruviana peristome absent, single costal stereid band, and hydric (Sull.) M. J. Cano & J. A. Jim´enez, of Bolivia, Chile, habitat. These four major traits are equivalent to four bits Ecuador, Peru, and Venezuela (Cano & Jim´enez, 2013), or 0.94 BPP of support for the distinction of Anoectan- and reexamination of the genus is needed at the world gium from the similar Streblotrichum,whichlikewise level. The genus Molendoa exhibits a clear reduction has sheathing perichaetial leaves. The fourth column 336 Annals of the Missouri Botanical Garden

of Table 1 shows traits of progenitor and descendants that evolutionary concepts are powerful decoding techniques are involved in evolution for this genus, not traits of because causality, even in the form of probabilistic dif- cladogram nodes. Nodes in cladistics do not model ferential survival, allows process-based explanations of evolution. Cladogram nodes are artifacts of the dichoto- organismal relationships. mous cladistic method of non-ultrametric hierarchic Classifications reflecting the new synthesis of adap- cluster analysis involving trait changes between species tation, population genetics, and development have been and whole groups of taxa, and are not inferentially challenged by patterns with strongly supported BPPs in responsible for continuity of a lineage. Total Shannon phylogenetic likelihood analyses. The BPPs, however, informational bits for the genus Anoectangium, as support assigned to a combination of process-based adaptation for the integrity of the genus, is 19 bits, or a statistical near and probabilistic drift in Table 1, are a middle ground. certainty for this branched lineage. Anoectangium is a Gould’s (2002: 893) speciational reformulation of closed causal group because speciation is operating at an macroevolution has a similar asymmetry and posits expected and acceptable three to five bits per event. This an evolutionary wall—in his example involving mini- count is about the same as the average four bits per mum structural constraints on size. For example, al- speciation event for Streptotrichaceae (Zander, 2018: 72, though there are mutations promoting variation toward 201) and five bits per event for Didymodon Hedw. both small and large size, there is a developmental (Zander, 2014: 16). Although some traits found useful limitation to small size for particular organisms, a kind by other authors are considered variable (leaf length, of phyletic constraint. He advanced a comparable ar- basal marginal crenulation, costal currency), this study is gument (Gould, 1970) regarding Dollo’s law rejecting able to avoid 1-character taxonomy, as shown in Table 1. complete reversal to a previous taxon state. An example The caulogram (Fig. 1) implied by the analytic key in systematics (e.g., as used in the present paper) is (Table 1) may be compared with the previous caulogram deciding that a complex, robust sexual species gave rise offered for the North American species of Anoectangium to very similar but reduced, simplified, asexual species (Zander, 2017a). It had the evolutionary formula A. because re-evolution of the lost traits is improbable. stracheyanum . ((A. sikkimense)(A. aestivum . A. One might argue that epigenetics may allow suppressed handelii)) with Hymenostylium xanthocarpum (Hook.) traits to be re-expressed, but that needs to be demon- Brid. as outgroup. At the time that study was done, A. strated on a case-by-case basis as that phenomenon has aestivum was considered a somewhat reduced derivative not been demonstrated as common and can be used to taxon, and the more robust A. stracheyanum was the best explain anything. choice of a generalist progenitor, while Molendoa anti- An amusing criticism one sometimes hears is that qua was mistaken for an Anoectangium. Presently, one’s rival scientist has shot an arrow, then painted an however, A. aestivum is viewed as a biotype-rich, wide- explanatory target around where it landed. This is hard spread progenitor, and A. stracheyanum is considered a to refute, mainly because it is an apt simile. The target is somewhat specialized descendant, though remaining a the relevant data set, the arrow is the method, and progenitor of A. sikkimense. The lesson here is that circles in the target gauge the optimal solution. In other macroevolutionary analysis is corrigible, with new, words, we impose a reasonable pattern on some aspect of process-based hypotheses developed based on new in- nature that fits the data as a theory or explanation with formation, and the new hypotheses are clearly advances predictive properties. Cladistics imposes a dichotomous that are directly explainable in terms of modification tree on data restricted to shared traits, which results with descent. in an optimal, well-supported but off-center target because it does not address information about ancestor- descendant relationships. In macroevolutionary system- DISCUSSION atics, we choose for analysis the target of a closed causal Wiley (1990), Brooks et al. (1986), and Brooks and group as best conceivable. Start with a group of max- Wiley (1988) have applied the concept of entropy to imally similar organisms assuming gradual steps in evolutionary analysis, and this may be extended to macroevolution. This is the essence of building on past systematics. In Shannon information theory (Shannon work by starting with the clustering of past classifica- & Weaver, 1949; Weaver, 1949; interpreted for sys- tions. Then we do or redo a homology analysis of our tematics by Zander, 2018: 31, 43, 47, 57), the highest chosen group. This decodes much of the information, entropy is equivalent to complete decoding of a cryptic i.e., increases informational entropy by recognizing message; the obverse, negentropy, is the potential or that a trait in one species is actually the same trait in power in a cryptic message to reveal information. Sys- another. Identification and elimination of species whose tematists seek to decode the hidden messages—the evolutionary processes involve sets of rather different negentropy—among the welter of information about traits purify the group. Subsequent analysis (the arrow) life in our natural world. Darwinian and neodarwinian by macroevolutionary evaluation using direct descent Volume 104, Number 2 Zander 337 2019 Macroevolutionary Evaluation with Anoectangium (Pottiaceae) in North America and the Himalayas increases entropy even more, being more efficient rather clearly evolutionarily redundant, in the sense of because the evolutionary explanation is simpler (no Zander (2018: 200 and following pages). It potentially postulated nodes or forced dichotomies). The use (ten- has the properties of survival during extreme climate tative imposition) of the theory of the dissilient genus change over hundreds of years, helping ensure a ther- concept, with taxa rather than nodes providing evolu- modynamically stable ecosystem on exposed wet cliffs. tionary continuity, is a better decoder than phyloge- It may serve as a point of origin of future biotypes and netics, which uses parsimonious dichotomous trees or perhaps species better adapted to changed environ- Markov chains, and shared characters of all distal ments. The fact that there is no evidence of sexual re- species (except reversals) gathered at dichotomous production seems no barrier to potential evolvability. For nodes as continuity. The optimum solution (the target’s instance, the asexually reproducing species Didymodon bull’s-eye) is the analytic key that minimizes reversals hedysariformis Otnyukova, D. murrayae Otnyukova, and and, after homology assessment and reevaluation of the D. zanderi Afonina & Ignatova all have demonstrably causal group, maximizes BPP. different infraspecific molecular races (Kuceraˇ & Ignatov, A Pleistocene species pump origin (Haffer, 2008; 2015), judging from sister grouping of the infraspecific Sedano & Burns, 2010; Schoville et al., 2012; Antonelli racial lineages. Simply speaking, homology assessment et al., 2015; Papadopoulu & Knowles, 2015) for North and closing the causal group removes “noise” and leaves American populations of Anoectangium sikkimense and only evolutionarily significant differences. A. stracheyanum may explain the broad spectrum of At this point one may note that the results of this study comparatively poorly differentiated populations of these and of my previous macroevolutionary analyses, although species in the British Columbia and Alaska region individually novel and significant, are part of an “exis- compared to strongly differentiated specimens exam- tence proof” that such analyses can successfully address ined from other areas of the New World. Repeated all data, allow predictive hypotheses, and provide well- isolation during glaciation is the “pump” that encour- supported evolutionary information about relationships ages biotype differentiation. The genus Anoectangium is that is more exact and direct than that of cladistic studies. old, judging from the lack of a peristome and reduction of the costal morphology, yet re-establishment in gla- ciated areas may not be easily explained by migration Literature Cited from refuges of presently rather stenomorphic A. aesti- Allen, B. 2002. Moss Flora of Central America. Part 2. vum and A. stracheyanum of North America to the east Encalyptaceae–Orthotrichaceae. Monogr. Syst. Bot. Mis- and south. souri Bot. Gard. 90. Allen, J. C., W. M. Schaffer & D. Rosko. 1993. Chaos reduces A more plausible theory is wind-mediated (by stormy species extinction by amplifying local population noise. westerlies) distribution of a set of fairly generalized and Nature (Taipei) 364: 229–232. polymorphic species from the unglaciated far northwest. Antonelli, A., A. Zizka, D. Silvestro, R. Scharn, B. Cascales- This takes the form of a species pump from a cache of Miñana & C. D. Bacon. 2015. 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