Whatever happened to Bishopanthus (Compositae, Liabeae)?

Abstract The enigmatic monospecific Bishopanthus of the tribe Liabeae (Compostiae/Asteraceae) has never been fully understood. It has not been possible to examine it in detail since it was described, because most of the type material was destroyed shortly after it arrived at the US National Herbarium, the morphology is insufficient to assign it to a subtribe, and the small amount of leaf material that remains is unsuitable for DNA extraction. A detailed description in English, image of the type specimen, photograph, and original illustration are included along with an estimation of where it was collected in the hopes that this information will encourage other field botanists who collect in northern Peru to search for it.


Introduction
Robert Merrill King (1930King ( -2007 was associated with the Smithsonian Institution from 1962 until 1998. During that time he organized many collecting trips to gather Compositae, mostly in the Neotropics. Among the most successful trips made by King, were those with Luther Earl Bishop (1943Bishop ( -1993. Th us, it was during a trip to Peru in early 1983 that Bishop, who frequently left the roadside and hiked into nearby areas, walked up a trail beside a small shop with hanging potted plants and returned within a half hour with a plant that has caused us problems for 30 years. Problem 1) Th e collection arrived at the US National Herbarium (Smithsonian Institution) only a short time before the publication of a monograph by Robinson (1983a) that was supposed to be a complete summary of the Asteraceous tribe Liabeae. Th e collection was immediately recognized as a new genus (by HR) and, in eff ect, it made the monograph he had completed obsolete before it was published. Bishopanthus was published later that year in Phytologia (Robinson 1983b).
Problem 2) Th e original fi ve ample sheets of the gathering were destroyed except for a few small scraps hidden under a bookcase that ultimately became the type.
Problem 3) Attempts by Michael Dillon (F) to collect the plant, based on information from King provided to Robinson, failed to fi nd any sign of the species.
Problem 4) Th e locality data were skimpy and the original collector, Bishop, and the leader of the expedition, King, have since died, so it is not possible to obtain any additional information.
Problem 5) Morphological analyses have failed to defi nitively place the species in a subtribe (Funk et al. 2012). Some of the characters link it to the Liabinae and others to a more remote position, as yet undefi ned. As a result, the genus is currently designated insertae sedis .
Problem 6) Dried leaf material that was collected for chemical analysis (Singh et al. 1985) was apparently discarded after the retirement of both Bohlmann & Jakupovic.
Problem 7) During the King & Bishop expeditions the herbarium material was placed in alcohol before being dried and so it is not suitable for current methods of DNA analysis.
In summary, what survives of the species is the herbarium specimen that has only the small scraps that became the type (Fig. 1), a color photograph taken by Earl Bishop (Fig. 2;Funk et al. 1996: 550, in black and white), a microscope slide prepared by Robinson from material before the specimens were destroyed (remade from the original Hoyer's solution slide during this study), the results of the chemical analysis by Singh et al. (1985) from a sample sent to him by R.M. King, and a picture of the pollen published in the pollen study of the Liabeae (Robinson and Marticorena 1986). From this material we have reconstructed the plant and an illustration has been produced (Fig. 3).

Materials and methods
Th e dried specimen material of Bishopanthus (what remained of the type specimen after it was destroyed) consisted of small fragments of leaves, a few small stems, an immature head and one mature head with three broken ray fl owers (Fig. 1). Th e disk fl owers faired better because they were protected if somewhat compressed. Additional material consisted of one slide of the dissected disk fl owers with a complete ray and a sectioned disk fl ower showing the anthers and style. From this material Tangerini was able to piece together the leaf fragments to form a few almost whole leaves and mounted these and other fragments and stems on two archival 4" by 5" cards ( Fig. 1). Tiny mylar packets were made to hold the two heads and dissected disk fl owers. Drawings of all of the fl oral details were made using a combination of the dried and slide material and the habit was reconstructed from the single photograph which provided an invaluable reference for the structure of the habit and leaf insertion. Conversations with Robinson (who had seen all of the original fi ve duplicates) provided information on the fl ower position on the stem. Th e fi nal drawing was done with a variety of polycarbonate pencils (Turquoise Filmograph, Mars Duralar and Mars Dynagraph, and L. & C. Hardtmuth Koh-I-Lar pencils) on Grafi x double matte drafting fi lm. Th ese pencils produce a denser black than regular graphite pencils and the subtle gray tones give the impression of the bullate leaf surface and the woolly texture of the involucres. Shading is applied with the softer leads fi rst followed by the harder leads to add fi ne lines and smaller details. Th e polycarbonate leads are waxier and do not smudge as much as graphite. Unfortunately the polycarbonate leads went out of production in the mid 1990's so this technique has a limited lifetime dependent on acquisition of the remaining stock of pencils. Some botanical art schools have supplies for illustration classes and perhaps resurgence in interest will spur new production.
Th e microscope slide was made using Hoyer's solution with a formula as given by Anderson (1954): Distilled water 50 cc; Gum arabic (U.S.P. Flake) 30 grams; Chloral hydrate 200 grams; Glycerin 20 cc. Since the original application of this method, Chloral hydrate has become a controlled substance, which limits access. More important is the impermanent nature of Hoyer's slides unless the cover slips are ringed with waterproof sealant (e.g., clear fi ngernail polish). Th e fact that microscope slides in Hoyer's solution can dry up does not mean they should be discarded. Th e water  miscible solution that makes the medium so useful as a rapid mounting medium also allows for recovery of any material on old dried slides. Th is should be noted by anyone inclined to discard old slides that often contain valuable type material.

Systematics
Diagnosis. Small shrub with milky sap; opposite leaves that are tri-nervate with a bullate upper surface and an under surface covered with a dense, grayish-wooly tomentum; two rows of subequal involucral bracts; short (~ 1mm) style branches; 8-10 ribs on the achenes and a bi-seriate pappus of subequal bristles (Table 1).
All members of the Liabeae have opposite leaves although some have very short internodes so they appear to have rosettes. Also, all members have milky sap and/or wooly light colored tomentum on the undersurface of the leaves.
Th e specimen label says that the plants are ½ m tall, have yellow fl owers and lots of milky sap. An examination of the fi eld notebook from the King & Bishop expedition did not provide any additional information about the plant or the location, however, an examination of their route that day shows that on the 21st of January their collecting started as they drove south from Tingo (6°22'18.14"S, 77°54'38.54"W). Th eir fi rst stop was 3 km south of Tingo and they proceeded south stopping occasionally until they reached their fi fth stop at 31 kms south of Tingo, probably near the village of Yerbabuena (6°33'57.92"S, 77°49'49.35W). Th e fi rst fi ve stops are all listed as ca. 5500 ft in elevation (based on Google Earth this seems a bit low) but with diff erent kilometer distances. Th e next stop (# 6) is where they collected Bishopanthus , however at this point they stopped giving the kilometers and just said "mountains behind Tingo ca. 6500 ft"and it is the only collection at that stop. Th e fi nal stop of the day (# 7) has the same location with an elevation of 7000 ft. At the 7 th stop they collected Cronquistianthus bishopii King & H. Rob., also restricted to this area. Th e next day (22 January 1983) the fi rst collecting stop was 61 kms along the road from Chachapoyas NW of Jaen. Evidently they were working out of Chachapoyas. We think that on the 21st of January they turned around at the 5th stop (31 km) and headed back to Chachapoyas, passed through Tingo, and stopped somewhere on the more eastern road between Tingo and Chachapoyas at 6500 Ft. Th e elevation on this road reaches 9000 ft. According to Google Earth the elevation is just about correct at the following coordinates: 6°21'40"S; 77°54'4"W. Of course this is a guess because the real location has never been found.

Conclusion
In their 2012 paper Funk et al. suggested that the shrubby habit and bullate leaves of Bishopanthus were similar to the taxa found in the basal grade of the subtribe Liabinae (e.g., Ferreyranthus ), but those genera lack latex and have pinnate venation and long style branches, in contrast to Bishopanthus (Table 1). Bishopanthus shares a number of characters with Cacosmia which is sometimes placed as the sister group of the Liabinae (Table 1). But Cacosmia has lost its pappus and has a unique (to Liabeae) achene (Table  1; fi rst placement). In addition, Cacosmia is unique in the tribe in that it has a small cylindrical head (5 ray fl owers and 5-6 disc fl owers) with a highly imbricate involucre (5 series and mostly ranked) while Bishopanthus has a broadly campanulate head (like most of the other genera in the tribe) that is larger (20/25) and an involucre of two subequal series; it also has a very diff erent habit. Now that we have the additional information on Bishopanthus , it looks as though it may be related to the Austroliabum element of Microliabum (Table 1; second placement) except that it does not have bullate leaves. Such a relationship would place it in the Paranepheliinae subtribe. Typical Austroliabum has been placed with some doubt as a synonym of Microliabum. Two of the typical species of Microliabum have been sequenced and now we need fresh material of Bishopanthus so we can see if either of our predictions is correct.